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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
2001. The second. For data acquisition the complete transect length of 600 m was subdivided in 25 m subunits (24 subunits/transect). The starting coordinate for each transect always marked the southeast corner to ensure identical geographic orientation between sampling units. four in secondary forest). A minor rainy season lasts from March/April to July. map “Projet OMS Forêt de Taï”). Richards 1996). More detailed descriptions of TNP are provided by Rompaey (1993). Doan 2003) or total removal plots (Barbault & Trefaut Rodrigues 1978. (1994) and PACPNT (2000). 5°50’N. PACPNT 2000). We thereby evaluated several methods that are already commonly used and present a set of techniques that. 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. Every subunit was marked with numbered colored flag-tape. Riezebos et al. known as transect sampling. on the basis of our experience in the field as well as with consecutive data analyses. Whereas quadrat sampling can be used to determine the species present in a homogenous area. situated in western Côte d’Ivoire (5°08’6°07’N. Floristically the TNP belongs to the Guinean-Congo-Region (Guillaumet 1967. The major rainy season stretches from September to October. We avoided extensive cutting and thus manipulation of important habitat features. Vonesh 2001. Given the lack of time for the development of effective conservation programs. known as quadrat sampling (plot design). 2001) were a priori excluded from testing. The rectangular transect design is a combination of two widely used standard techniques. that ii) is easy to handle. A number of recently published regional field monitoring manuals for amphibian surveys (Latin America: Lips et al. b. thus ensuring that all habitat types of a certain area within the very inhomogeneous forest were covered.RÖDEL & ERNST essential for academic and practical purposes alike. 7°20’W). also for para-ecologists. 6°47’-7°25’W). standardization of methods that yield broad-scale comparative data is now needed more than ever. uses linear transects instead of squares. and iv) has a low environmental impact. Precipitation is distributed across two more or less distinct periods. that stretches west of the Dahomey Gap from Ghana into Sierra Leone and Guinea (Schiøtz 1967. It can be characterized as humid tropic seasonal (Riezebos et al. data: Taï Monkey Project). 1979a. 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. 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. Parts of the habitats on the park’s periphery were subject to more or less intense logging and cultivation until 1998 (P. such as standard plot sampling (Allmon 1991. Daily tempe2 . We aim to present a methodology that i) provides quantitative and qualitative data useful for rapid surveys as well as long-term monitoring programs. consists of a series of small squares (quadrats) that are laid out at randomly selected sites within a habitat then thoroughly searched for amphibians. and unpubl. All evaluations and recommendations are based on a study of tropical amphibian assemblages conducted over four years (1999–2002) in Taï National Park. The maximum distance between transects was 6. Each had a north-south extension of 200 m and an east-west extension of 100 m. 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. Rodda et al. Parren & de Graaf 1995. The first. Africa: Howell 2002) reflect this need.3 km. ratures vary between 20 and 33° C. 1994. as well as their relative abundances and densities. The core dry season lasts from November until February/March. Our study sites are either pristine forest areas or have not been logged or cultivated since 1978. The minimum distance between adjacent transects was 200 m. TNP is the largest remaining protected area of rain forest in West Africa. Lawson 1986. as they may result in severe disturbance to the system under investigation. Poynton 1999). transect sampling can provide similar data across habitat or dis- MATERIAL AND METHODS Study site The study was conducted in Taï National Park (TNP). TNP is part of the Upper Guinea forest block. The mean annual temperature is about 25° C (Rompaey 1993). followed by a short dry season in July/August. Mean annual precipitation in the study area was 1806 mm during 1988–2002 (± 297mm. Formenty pers. Thus widely used techniques. Transect paths were kept open so that walking at a constant speed was possible at all times. proved to be most effective for forest amphibians. iii) is efficient concerning time and money. Côte d’Ivoire. Transects were arranged in pairs. comm.
divided into seven categories corresponding to particular densities.g. These parameters included vegetation density in four strata (canopy: > 20 m. Sampling was performed independent of prevailing weather conditions. vegetation density was measured in four strata (see text). In order to quantify the availability of potential aquatic breeding sites. 1996. both in time and space. substrate moisture was determined in four categories during every transect walk. Repeated controls of identical transects on consecutive days were avoided to ensure independence of samples. Usually four to six transects per day were sampled during daytime. Compared to linear transects. Jaeger & Inger 1994). To avoid duplicate recordings.AMPHIBIAN MONITORING IN TROPICAL FORESTS turbance gradients (Jaeger 1994. for comparison of local with regional species pools). secondary forests. Categories of habitat variables measured on transects. whereas primary for- ests show increasing numbers of plants of larger dbh (Chatelain et al. to assign them to particular guilds). In addition to species identity we recorded sex and snout-vent-length (SVL) of every individual. lower tree stratum: 3–10 m. 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. Definitions of habitat variables are summarized in Table 1. 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. Sampling methods and sampling effort Habitat characterization.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 .g. bush and shrub stratum: 0.5 3 3. We assumed that the number of plants with small dbh is greater in degraded. and to investigate correlations of species assemblages with environmental variables..5 m).5 2 2.35 m/s). In addition. captured frogs were marked by TABLE 1.30–0. understory: < 0.5–1. Standardized visual transect sampling (SVTS). dbh).or age-specific differences in habitat use. Additionally. thus providing data that are useful to test for sex. Specimens heard but not seen within this distance were not searched for and not included in the SVTS data sets (see below). A maximum of four transects was patrolled at night. thereby recording all amphibians within a distance of 100 cm from either side of the path.. we characterized all habitats using several variables that were recorded at two defined points within each 25 m subunit (beginning and midpoint). 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). Transects were intensively patrolled at a constant speed (0. dbh = diameter of plants at breast height. To assess habitat preferences of particular species (e. Edaphic parameters were registered as general substrate types according to Lieberoth (1982). 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. Pearman 1997). As far as possible all individuals were captured. Sampling was interrupted for the duration of the recording in order not to overestimate locations in which animals had previously been captured. rectangular ones provide the possibility of easily up.or downscaling data without neglecting local habitat diversity (e. surface and depth.5 m.
0. Thus. Drift fences consisted of durable green plastic gauze. despite their potential abundance.g. Due to the simplicity of the method. Likewise microhabitat description was not always possible. 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). only calls coming from the right-hand side in the first segments were registered (i. As opposed to visual sampling..g. 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). VES and AES thus could be used to evaluate if the randomly chosen transects represented the regional species pool. beginning with segment 1: segment 1. 8.5 m high and stapled vertically onto wooden stakes. and to estimate relative abundances of species within an assemblage (Crump & Scott 1994). Ptychadena species) are more difficult to trap than terrestrial species that lack these abilities (e. For practical reasons “man-hours” can be used. An array of fences and traps consisted of a central trap (buckets: . 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. Visual (VES) and acoustic (AES) encounter surveys. species composition. 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. Dodd 1991). other than sex and species. 1997). Audio strip transects represent a commonly used method for acoustic monitoring (Zimmerman 1994). Since in the majority of frog species males use species-specific calls to advertise their position to potential mates and rivals (Wells 1977)..5 4 hours of transect sampling. Bufo species). This technique has been formalized as the time-constrained technique of Campbell & Christman (1982). It can be used to determine the species richness of an area and the species composition of a local assemblage. Throughout transect walks a combination of visual and acoustic techniques was applied at all times. 1994. In the transect corners. habitat analyses can be based on data obtained from the general habitat characterization for single transects or segments. this technique allows the detecting of cryptic species that. during SATS it was often impossible to determine individual parameters. Counts can be used to estimate relative abundance of calling males. Pitfall traps with drift fences were installed in two of the primary forest transects from March to April 1999. individual marking should be restricted to studies in which individual recognition is indispensable. This corresponds to 765 transect walks. 12 and 20). Recaptures were excluded from the analyses. Calls from greater distances cannot be unambiguously identified and the chance of duplicate recordings in neighboring segments increases. Furthermore. For that reason a maximum recording distance of 12. VES can only provide information on the presence or absence of a species in an area but are inadequate for determining abundances. For both SVTS and SATS. Total VES. as well as breeding habitat use and breeding phenology of species (Zimmerman 1994). Standardized acoustic transect sampling (SATS). and additionally in only one of these transects from January 2001 to September 2002. Henle et al.RÖDEL & ERNST toe clipping (Donnelly et al. VES and AES are frequently used for rapid assessments and the evaluation of larger areas. keeping the transect routine identical for both techniques. The resulting data are expressed in numbers of individuals of a certain species found in an area per unit time.and AES-effort was kept comparable to the time spent on SVTS and SATS surveys throughout the entire study period. From February 1999 to December 2000 we compiled 382. following the characterization routine used for general habitat description. this behavior can be exploited for acoustic monitoring. According to Corn & Bury (1990). Trapping with pitfall or funnel traps along drift fences. Coding schemes used for individual recognition were not applied. Anurans that are strong jumpers (e. which can be adjusted to the complexity of the habitats being sampled. In those cases. An area or habitat is searched systematically for individuals in a defined time period. This device is useful to determine species richness of epigaeic organisms (Corn 1994). Capture success may vary greatly between species (Corn & Bury 1990.e.5 m to either side of the transect was defined. Individuals below nine mm SVL were not marked due to their small size. may be underestimated when exclusively using visual techniques. data can be expressed in numbers of individuals per time and surface units. In concordance with these authors the methods were only used as a qualitative or semi-quantitative tool within the scope of our studies. thus creating 25 m x 25 m acoustic sampling plots.
using a particular sampling method. Sampling success is referred to as number of species within families recorded. arboreal. The ends of each segment were flanked with additional plastic buckets. They were checked at least every morning. SVTS = species recorded during visual transect walks. fossorial and aquatic species independent of families. Pitfall trapping time summed to a total of approximately 4000 trap-days. TC = species recorded during transect walks (SVTS and SATS combined). results were analyzed for leaf litter. TaïF = all species recorded with all methods in forest habitats around SRET station. 220 mm bottom diameter) and two triangular fence segments (total length 16 m). one on either side. 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. Hillers et al. AES/VES = visual and acoustic encounter surveys (only species that have been recorded in forest habitats around SRET station). Additionally. Each segment was tightened around a plastic bucket with an opening angle of 45°.AMPHIBIAN MONITORING IN TROPICAL FORESTS 275 mm deep. data). In the course of this study. 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). as sampling success and efficiency may vary with regard to the particular biology of a species. 1. Numbers above bars represent total number of species recorded with the respective method. Branch & Rödel 2003). 285 mm top diameter. funnel traps (see Branch & Rödel 2003) were installed in a primary forest transect from 18 to 27 September 2002. Trap = pitfall and funnel traps. Number of species per family recorded using different sampling methods.. The data from pitfall and funnel traps can be expressed as number of individuals per trap-day. unpubl. FIG. Traps were checked at least on a daily basis. we applied that method during a herpetological survey of the Haute Dodo and Cavally forests. Funnel traps summed to a total of 384 trap-days. 5 Ta ïF . which are situated south and west of TNP respectively (Rödel & Branch 2002. and in a forest fragment (Paulé-Oula 2) outside TNP from 30 August to 12 September 2002 (A. In addition.
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. . two species were each listed in two guilds (compare Appendix). Trap). Phlyctimantis boulengeri ). nienokouensis..g. Rödel 2000.RÖDEL & ERNST FIG. Sampling success and data quality varied between different methods.0 % of TaïF) in eight families. Seven species (Bufo maculatus.g. Number of species per amphibian guild recorded using different sampling methods. and P. Ten species (23..g. SVTS.2 %) were recorded with pitfall and funnel trapping. 2002. During transect sampling (SVTS. 2003. bibroni. Appendix 1). compare Appendix 1). VES/AES revealed 39 forest species (90. Ptychadena pumilio. Bufo superciliaris) or simply absence in the area under investigation (e. P. clearing around SRET) and thus ignored in the analysis. sp. Phrynobatrachus calcaratus. 1. annulatus) were exclusively recorded during transect walks. Rödel et al. Using all methods we recorded 50 species in the region of the SRET (89. Rödel & Ernst 2000. Hoplobatrachus occipitalis. tested for TaïF. 43 species were recorded in forest habitats within the study region (Fig. AES/VES. regularis. and between different amphibian families or guilds (Figs. Two fossorial species were exclusively detected either with VES and pitfall traps (Hemisus sp. SATS. see Appendix 1). 2. 2.) or pitfalls only (Geotrypetes seraphini). 1. H. Effectiveness in detecting species within given families differed significantly between methods (Friedman test. fusciventris. H.3 % of TNP species).) were recorded exclusively outside forest habitats (e. P. Families containing only 6 a few species in the TNP area showed the highest recording rates. 2). rather than actual sampling method insufficiency. For abbreviations see Fig. P. Perret 1988. B. N = 10. Phrynobatrachus fraterculus. Hyperolius guttulatus). TC.002. 1. Four arboreal frogs were exclusively recorded with these methods (Hyperolius picturatus. p = 0. Taking all methods into account. 56 amphibian species are known to occur in TNP (Schiøtz 1967. SATS) we recorded a total of 15007 individual amphibians belonging to 37 species (86.7 %) of nine families. Failure to detect particular species in this region was most probably due to their general scarcity (e.199. RESULTS In total. χ2 = 19. mascareniensis. Three leaf litter species (Amnirana occidentalis.. thus making overall sampling success very high (Tab. df = 5.
quantitative data for all arboreal species were best assembled with SATS.0 100. No additional species were detected with funnel traps. As aquatic sites were only taken into account. was difficult as detectability of species varied considerably with (e.5 100.7 0..0 50. in a primary forest transect with none in the forest fragment (Hillers et al. but very low in arboreal species (6. Most specimens were captured with funnel traps (0. Schaich & Hamerle post hoc multiple comparisons revealed no significant differences between methods at α = 0.5 % of recorded leaf litter frogs).0 72.0 100.0 100.0 AES/VES 0. SATS.0 0. other than based on time units.8 0.0 100.0 0.0 36.0 100. A distance of 1 m (visual) and 12. 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 0.0 100. We only captured a few leaf litter frogs. In contrast.0 100.0 100.0 31.294.0 100. transect walks made it possible to search for amphibians in a very comparative way.4 0.0 100.0 100.0 50. unpubl.0 77. Most arboreal frogs could be recorded with SVTS.0 64. data). 7 .0 100.7 100.0 76.) vegetation density.067.0 76. which is mute.0 100. all were recorded using other methods as well. For other abbreviations see text and Fig.0 100.8 0. neither between given family or guild.7 % of recorded arboreal species). but that concerned mostly single specimens. As in TNP.AMPHIBIAN MONITORING IN TROPICAL FORESTS TABLE 2.5 m (acoustic) proved to work equally well in all habitats investigated. when being a part of the transect line itself. However.0 100.0 SATS 0. All other species were recorded with VES/AES (Rödel & Branch 2002. p = 0. TC.0 100.0 100.0 86.0 100.0 0.0 Trap 100. Trapping success was highest in leaf litter frogs (56.0 66. Percentage of species within an amphibian family or guild that were recorded using a particular method. χ2 = 10.5 100.05.0 100.03 specimens per trap-day in pitfall traps). SVTS.g. the fossorial Geotrypetes seraphini was recorded with pitfall traps only.0 0.0 100. this was most probably due to the small overall sample size and high heterogeneity of species numbers in families and guilds (Bortz et al.0 100.0 77. 1990). Trap). 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). 1. N = 4.0 81.4 100.0 100.7 0.8 0.2 66.0 0. Likewise.47 specimens per trap-day in contrast to 0. VES and AES revealed highest species numbers.0 0.8 100. presence and abundance of purely aquatic species and tadpoles of all species were underestimated with all methods used here.0 0.7 100. In TNP funnel traps were ineffective. Branch & Rödel 2003). asytlosternids) were best recorded with SATS.0 100.0 100. df = 4. Diurnal ground-dwelling frogs were best recorded quantitatively using SVTS.0 33. nocturnal leaf litter frogs (arthloeptids.3 We could not detect a statistical difference in sampling efficiency between different amphibian guilds using the methods employed (Friedman test. With the exception of Acanthixalus sonjae.0 100.0 40. 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. tested for TaïF. Quantification of these data.0 100.0 0.0 80.0 68.0 SVTS 0. VES/AES. Reference is the number of species that were recorded using all methods in forested habitats around the SRET station (TaïF).0 100.
In this study this has been done by determining relative species richness with reference to families or guilds. were nearly as successful in detecting species. However. We found it more realistic to apply a (clearly subjective) measure for habitat complexity on which to base collection effort during VES/AES. The methods VES and AES in combination with pitfall traps with drift fences were useful in sampling additional. (1995) a simple comparison of species numbers may not be sufficient to evaluate the efficiency of different sampling methods. spending more time in complex. especially in long-term field studies within temperate regions (e. Cardioglossa leucomystax. The methods tested within this study proved to perform with varying degrees of efficiency. could be detected exclusively using visual sampling methods.RÖDEL & ERNST DISCUSSION Choice of methods should reflect the best compromise between i: practicability. When including relative abundance measurements these methods were insufficient. this only holds true if species numbers alone are considered. Doan 2003). ii: specific aims.. They suggest using relative species richness within taxonomic entities.1. as well as SVTS alone. Methods of collecting quantitative data for aquatic amphibians and for tadpole assemblages have been described by Shaffer et al. Semlitsch et al. and will likewise hold true for other arboreal anurans throughout the tropics. such as the very abundant Arthroleptis sp. a combination of SVTS and SATS. We found it impossible to quantify VES and AES other than in relation to time. Whereas transect walks provide the possibility of monitoring amphibians in a very comparative way. whereas visual sampling appeared to be superior to acoustic sampling. Thus even time-based quantification of these data is questionable. VES and AES are useful tools for the compilation of species inventory lists.2. such as species of the genus Acanthixalus. Arthroleptis sp. 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. Although the combination of AES and VES provided an adequate approximation to real conditions (see also Doan 2003). such as species groups. The combined transect sampling methods provided a close approximation to the real presence and abundance of species. especially arboreal and fossorial. data acquisition varies considerably depending on habitat type in VES and even AES. Simple presence/absence data of aquatic species and stages can best be gathered by direct observation (surfacing individuals) and with dip-netting. 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. (1994). Nonetheless. Migrating pipid frogs can also be easily recorded with pitfall and funnel traps. 2003).g. genera or families. SVTS was generally the best method for detecting diurnal leaf litter frogs. (1997) and Rödel (1998a). in order to sample particularly secretive leaf litter species. In general SATS was very efficient in detecting nocturnal leaf litter frogs and treefrogs but less successful in detecting diurnal leaf litter frogs. during rapid assessment surveys (for recent West African examples see Rödel & Branch 2002. Rödel et al. Vonesh 2001. Pitfall trapping was only useful in detecting fossorial species that were not encountered during transect walks. and iv: environmental impact. We found no additional species and only very few specimens by funnel trapping. such as the Hylidae. these methods do not seem to sample a given fauna adequately well (Allmon 1991. Schiøtz 1999. as they impose a severe disturbance on the system under investigation and thus were not suitable to our goals. VES and AES yielded highest species numbers. Standard plot sampling and total removal plots were not tested. e. According to Pearman et al. iii: efficiency. determined by using a particular sampling method. This was also true for most of the arboreal species belonging to the families Hyperoliidae and Rhacophoridae. Olson et al. Although widely used and recommended. and failed to detect aquatic and fossorial species. as an adequate measure. can then be compared to the real conditions in an area or habitat. The percentage of species within supraspecific taxonomic units. species not encountered during transect sampling. The differences in efficiency of visual versus acoustic sampling varied between taxa. 1996). inaccessible areas than in areas that are easy to monitor. Rödel 2003. pitfall traps and drift fences . However. or Phrynobatrachus alticola. The advantage of transect sampling was especially obvious when focusing on leaf litter species. Some taxa. In addition. acoustic sampling appeared to be indispensable since these species were more readily detected by their calls than 8 by sight. but is certainly just as well suited for the investigation of tropical anuran assemblages in general. which are mute (Drewes 1984. Rödel & Ernst 2003).. Bury & Corn 1987.g.
Parris 1999. in consecutive field seasons. Burger et al. Donnelly et al. experience of investigator and prevailing weather conditions (see Branch & Rödel 2003). as well as environmental changes. Rodda et al. VES and AES are less time consuming. Traps have to be checked at least daily. However only two species. When considering the relatively high costs and time investment due to the high level of maintenance required (Parris 1999). one should test the efficiency at a particular site in advance before installing these traps on a broad scale. in theory. amphibian guild and environmental impact. 1.) that leopard (Panthera pardus) tracks TABLE 3. Doan 2003). However. A comparison of trapping data with data other than those of the observer in question seems to be difficult. SVTS and SATS have been proven to be the only methods that provided quantitative data on forest amphibians with regard to space and time. Transect walks are time intensive. the fossorial Hemisus perreti and the aquatic Silurana epitropicalis. They collected only six species (out of a total of 132 species of amphibians and reptiles being recorded). this method cannot be recommended for standardized sampling of tropical anurans. 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. (2001) report on similar experiences using this method in a herpetological survey in Guyana. Theoretically it might well be that trapping will result in dislocating at least some species from the area where traps have been established. Data from both trapping methods can. The impact of traps and drift fences was difficult to judge. seasons and years. 2001. We observed (e. traps have to be uninstalled or closed and reopened when starting again. however. 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 . (2004) had high trapping success with pitfalls in Gabon when considering numbers of individuals. However trapping success depends very much on choice of site. Data can be used for comparisons between habitats.AMPHIBIAN MONITORING IN TROPICAL FORESTS proved to be the least effective method. Costs with regard to time spent for data acquisition was highest in trapping and in transect walks. It is advisable to perform them randomly independent of prevailing weather conditions. If the investigation is interrupted for some time. QS/TR = quadrat sampling/total removal sampling (data from Allmon 1991. for other abbreviations see text and Fig. none of them uniquely found by this method. VES and AES had probably the lowest environmental impact. Effectiveness of methodology in regard to type of data required. Transects will have an environmental impact. Vonesh 2001. accounted for 72.g. e. if not on amphibians then on other organisms. SVTS and SATS provided data on species and habitats und thus may reveal changes in composition and species abundance. In our experience. as trapping success per trap per time unit.6 % of all individuals captured with this method. be quantified in relation to time. generalization of data with regard to phenology or abundances is less possible..g.
R.A.D. TROPENBOS . & K. (ed. R. Furthermore transects provide an effective method of investigating leaf litter frogs in particular. (eds. Sci. ACKNOWLEDGMENTS This study is part of the BIOLOG-program of the German Ministry of Education and Science (BMBF. J.. Transect cutting and the use of transects should therefore be done with the necessary precautions.” G. A. Trefaut Rodrigues. Herpetological survey of the Haute Dodo and Cavally forests. W. & A.. Wildlife Management 51: 112–119. Hammond. 01 LC0017). J. Global amphibian declines: a problem in applied ecology. SVTS and SATS are especially useful for long-term studies. After using transects intensively for some time. Pflanzensoziologie.Y. 939 pp. Memoir 28: 145–186.): Measuring and monitoring biological diversity. Trop. N’Dri. REFERENCES Adis. McDiarmid. Inst. J. Verteilungsfreie Methoden der Biostatistik.. Evaluation of pitfall trapping in northwestern forests: trap arrays with drift fences. Gautier. Pp. Sér. Basset. Trefaut Rodrigues.. Smithsonian Institution Press. Christman. Biodiv. in the worst case.Côte d’Ivoire helped with transportation. Bortz. Calif. N.” of the Republic of Ivory Coast. W.S.. A plot study of forest floor litter frogs. Field techniques for herpetofaunal community analysis..R. Springer Verlag. for specific questions (cf. U.. Observations sur la reproduction et la dynamique des populations de quelques anoures tropicaux I. & S.1964. A recent history of forest fragmentation in southwestern Ivory Coast. & S.. Branch.. Canopy fogging of an overstory tree – recommendations for standardization. 10 . Salamandra 39: 91–110. W. Donnelly. Ecotropica 4: 93–97. & M. Phrynobatrachus accraensis. & R. Allmon. All this support is gratefully acknowledged. Halliday.-O. P.P. Lips et al. Observations sur la reproduction et la dynamique des populations de quelques anoures tropicaux III. Central Amazon. M. a well visible path becomes established that at least in primary forests remains so for years. However. 1982. R.. 1979a. 20: 64–77. Pp. Terre Vie 32: 441–452. Foster. Conserv. Spichiger.W. Branch. and C. Barbault. & M. A 41: 417–428.B. Syst. Wildlife Research Report 13. R. 1998. Ecol. Gabon: Species turnover along an elevational gradient. J. 30: 133–165. 193–200 in Scott. & M. & M. C. Ouoro provided invaluable help during fieldwork.G.” Working in TNP was kindly permitted by the Commandant K. 1987. J. Advantages and disadvantages of different methods are summarized in Tab. consequent long-term effects on forest structure.-A. Rev. Jr. Floren. U. fond. T. Gbamlin. M. 1991. P. Springer Verlag. 1994. 1994. Research permission was given by the “Ministère de l’Enseignement Supérieur et de la Recherche Scientifique..A.. Fish and Wildlife Service 13. Observations sur la reproduction et la dynamique des populations de quelques anoures tropicaux IV.): Herpetological communities. Braun-Blanquet. Brazil. Glos. thus generating a much more complete picture of an amphibian community than is possible with other methods....R. 1990. 1997. 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.. Arthroleptis poecilonotus. Washington & London. 1979b. 1978. The “Projet Autonome pour la Conservation du Parc National de Taï”. Hillcoat provided valuable comments on the manuscript. Ptychadena maccarthyensis et Ptychadena oxyrhynchus. Linsenmair. Richards. Acad.. Part II: Trapping results and reptiles.J. & K.R.. Campbell. 1996. 2003) transects should be supplemented by specifically investigating aquatic breeding sites with the appropriate set of standardized techniques (Heyer et al. Department of the Interior. This might affect densities of other mammals including possible seed dispersers with. possibly accompanied by opportunistic trapping.S. Corn. The access permit to TNP was issued by the “Ministère de la Construction et de l’Environnement. Olson et al. Afr. Y. Lienert. 2 ed. western Ivory Coast.. Barbault.S. 3. Bull. 364 pp. Burger.W. Hofer. R. & M. 109–117 in Heyer. Boehnke. 5: 37–53. noire. Trefaut Rodrigues.RÖDEL & ERNST were more often seen on the transect path than in undisturbed forest. 7: 503–522. L.M. Ecol.A. Bury. Rödel 1998a). and the “Taï Monkey Project” provided logistic support.C. Barbault. and B.o.J. 2004.. L. 2003. Straight-line drift fences and pitfall traps.E. Project W08 BIOTA-West. Rödel. Amphibians and reptiles of Monts Doudou. & P. 1999.. J. Hayek. Trop. 865 pp. Alford. Channing. Ann. Chatelain. 1964. H. Grundzüge der Vegetationskunde. not only on their breeding sites but throughout the whole range of habitats used by them. The presented transect design proved to be adequate for representative sampling and appeared to be appropriate for most studies involving multivariate structure data. Standard methods for amphibians.S. W. Ecol. Corn. G. For simple short-term surveys we recommend VES and AES. Berlin a. R.. Wien..
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f = fossorial species. a = arboreal species. 2003). Perret 1988) but not recorded by us. 1 = known from TNP (Schiøtz 1967. 5°25’ N. G = species recorded in southern TNP (Guiroutou. TC = all species recorded on transects. 7°10’ W). 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. List of amphibian species recorded in Taï National Park (TNP). TS = all species recorded on transects in secondary forest. 4 = Acanthixalus sonjae lives aquatically in large water filled tree holes but forages in surrounding vegetation (Rödel et al. 2 Ranidae Hoplobatrachus occipitalis Amnirana albolabris Amnirana occidentalis Aubria occidentalis Ptychadena pumilio Ptychadena bibroni Ptychadena mascareniensis Ptychadena superciliaris Ptychadena aequiplicata Ptychadena longirostris Ptychadena sp. Trap = species recorded with pitfall and funnel traps. l = leaf litter species. 3 = Phrynobatrachus guineensis is a leaf litter frog but reproduces in small water-filled tree holes (Rödel 1998b). species Gymnophiona Caecilidae Geotrypetes seraphini Anura Pipidae Silurana tropicalis Bufonidae Bufo regularis Bufo maculatus Bufo togoensis Bufo taiensis Bufo superciliaris Hemisotidae Hemisus sp. a l l l l l 13 . SRET = species recorded with VES/AES on the clearing around the SRET station. aq = aquatic species. 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. 5 = treated as full species and not as subspecies of Hyperolius fusciventris due to sympatric occurrence in TNP (see Rödel & Branch 2002). 2 = determination not possible at present time. 6 = recorded north of area under investigation. TP = all species recorded on transects in primary forest. other than transects.AMPHIBIAN MONITORING IN TROPICAL FORESTS APPENDIX.
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. 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 .
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