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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
Riezebos et al. Given the lack of time for the development of effective conservation programs. For data acquisition the complete transect length of 600 m was subdivided in 25 m subunits (24 subunits/transect). A number of recently published regional field monitoring manuals for amphibian surveys (Latin America: Lips et al. Doan 2003) or total removal plots (Barbault & Trefaut Rodrigues 1978. situated in western Côte d’Ivoire (5°08’6°07’N. Daily tempe2 . The rectangular transect design is a combination of two widely used standard techniques. (1994) and PACPNT (2000). The first. 2001. b. Vonesh 2001. Whereas quadrat sampling can be used to determine the species present in a homogenous area. Parren & de Graaf 1995. as well as their relative abundances and densities. The core dry season lasts from November until February/March. The maximum distance between transects was 6. Every subunit was marked with numbered colored flag-tape. ratures vary between 20 and 33° C. Africa: Howell 2002) reflect this need.3 km. It can be characterized as humid tropic seasonal (Riezebos et al. The major rainy season stretches from September to October. Floristically the TNP belongs to the Guinean-Congo-Region (Guillaumet 1967. Thus widely used techniques.RÖDEL & ERNST essential for academic and practical purposes alike. Each had a north-south extension of 200 m and an east-west extension of 100 m. TNP is part of the Upper Guinea forest block. Mean annual precipitation in the study area was 1806 mm during 1988–2002 (± 297mm. TNP is the largest remaining protected area of rain forest in West Africa. thus ensuring that all habitat types of a certain area within the very inhomogeneous forest were covered. uses linear transects instead of squares. followed by a short dry season in July/August. Richards 1996). We aim to present a methodology that i) provides quantitative and qualitative data useful for rapid surveys as well as long-term monitoring programs. and iv) has a low environmental impact. Parts of the habitats on the park’s periphery were subject to more or less intense logging and cultivation until 1998 (P. 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. as they may result in severe disturbance to the system under investigation. proved to be most effective for forest amphibians. transect sampling can provide similar data across habitat or dis- MATERIAL AND METHODS Study site The study was conducted in Taï National Park (TNP). data: Taï Monkey Project). that stretches west of the Dahomey Gap from Ghana into Sierra Leone and Guinea (Schiøtz 1967. 2001) were a priori excluded from testing. PACPNT 2000). Precipitation is distributed across two more or less distinct periods. 5°50’N. The second. 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. 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. A minor rainy season lasts from March/April to July. also for para-ecologists. on the basis of our experience in the field as well as with consecutive data analyses. Formenty pers. 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. comm. four in secondary forest). Transects were arranged in pairs. known as transect sampling. The starting coordinate for each transect always marked the southeast corner to ensure identical geographic orientation between sampling units. Lawson 1986. standardization of methods that yield broad-scale comparative data is now needed more than ever. We avoided extensive cutting and thus manipulation of important habitat features. iii) is efficient concerning time and money. 6°47’-7°25’W). 7°20’W). The mean annual temperature is about 25° C (Rompaey 1993). Transect paths were kept open so that walking at a constant speed was possible at all times. that ii) is easy to handle. such as standard plot sampling (Allmon 1991. Rodda et al. 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. and unpubl. map “Projet OMS Forêt de Taï”). 1994. 1979a. Our study sites are either pristine forest areas or have not been logged or cultivated since 1978. Poynton 1999). More detailed descriptions of TNP are provided by Rompaey (1993). Côte d’Ivoire. consists of a series of small squares (quadrats) that are laid out at randomly selected sites within a habitat then thoroughly searched for amphibians. known as quadrat sampling (plot design). The minimum distance between adjacent transects was 200 m.
Categories of habitat variables measured on transects.. lower tree stratum: 3–10 m. and to investigate correlations of species assemblages with environmental variables.5 2 2. vegetation density was measured in four strata (see text). for comparison of local with regional species pools). definition category 1 1.5 m. 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. Additionally. These parameters included vegetation density in four strata (canopy: > 20 m.30–0. Sampling was interrupted for the duration of the recording in order not to overestimate locations in which animals had previously been captured.or downscaling data without neglecting local habitat diversity (e. As far as possible all individuals were captured. Transects were intensively patrolled at a constant speed (0. to assign them to particular guilds). In addition to species identity we recorded sex and snout-vent-length (SVL) of every individual. To avoid duplicate recordings.AMPHIBIAN MONITORING IN TROPICAL FORESTS turbance gradients (Jaeger 1994. thus providing data that are useful to test for sex. Edaphic parameters were registered as general substrate types according to Lieberoth (1982). Standardized visual transect sampling (SVTS). Specimens heard but not seen within this distance were not searched for and not included in the SVTS data sets (see below). Pearman 1997). divided into seven categories corresponding to particular densities. substrate moisture was determined in four categories during every transect walk.35 m/s). captured frogs were marked by TABLE 1.g. 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. 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. Repeated controls of identical transects on consecutive days were avoided to ensure independence of samples. Sampling was performed independent of prevailing weather conditions. We assumed that the number of plants with small dbh is greater in degraded.g.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 . A maximum of four transects was patrolled at night. To assess habitat preferences of particular species (e. dbh = diameter of plants at breast height. rectangular ones provide the possibility of easily up. both in time and space.or age-specific differences in habitat use. Definitions of habitat variables are summarized in Table 1. whereas primary for- ests show increasing numbers of plants of larger dbh (Chatelain et al. Sampling methods and sampling effort Habitat characterization.5 3 3. secondary forests. Jaeger & Inger 1994). we characterized all habitats using several variables that were recorded at two defined points within each 25 m subunit (beginning and midpoint).. 1996. In order to quantify the availability of potential aquatic breeding sites. surface and depth. thereby recording all amphibians within a distance of 100 cm from either side of the path. In addition. Compared to linear transects. dbh). Usually four to six transects per day were sampled during daytime.5–1. understory: < 0.5 m). For a discussion of the advantages of rectangular sample units over quadratic or circular ones see Krebs (1989) and McCune & Grace (2002). bush and shrub stratum: 0.
It can be used to determine the species richness of an area and the species composition of a local assemblage. Due to the simplicity of the method. VES can only provide information on the presence or absence of a species in an area but are inadequate for determining abundances. We used these methods continuously in all habitats of the study area including the transect areas. beginning with segment 1: segment 1.. and as the time-constrained searches of Corn & Bury (1990). Individuals below nine mm SVL were not marked due to their small size. Calls from greater distances cannot be unambiguously identified and the chance of duplicate recordings in neighboring segments increases. thus creating 25 m x 25 m acoustic sampling plots. 8.and AES-effort was kept comparable to the time spent on SVTS and SATS surveys throughout the entire study period. Total VES. 12 and 20).e. 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. Audio strip transects represent a commonly used method for acoustic monitoring (Zimmerman 1994).. individual marking should be restricted to studies in which individual recognition is indispensable. Likewise microhabitat description was not always possible. habitat analyses can be based on data obtained from the general habitat characterization for single transects or segments. as well as breeding habitat use and breeding phenology of species (Zimmerman 1994). Recaptures were excluded from the analyses. species composition. Furthermore. Bufo species). 0. 1994.. In concordance with these authors the methods were only used as a qualitative or semi-quantitative tool within the scope of our studies. may be underestimated when exclusively using visual techniques. this technique allows the detecting of cryptic species that. Throughout transect walks a combination of visual and acoustic techniques was applied at all times. Henle et al. According to Corn & Bury (1990). which can be adjusted to the complexity of the habitats being sampled. Counts can be used to estimate relative abundance of calling males. following the characterization routine used for general habitat description.5 m to either side of the transect was defined. and to estimate relative abundances of species within an assemblage (Crump & Scott 1994). As opposed to visual sampling. and additionally in only one of these transects from January 2001 to September 2002. Pitfall traps with drift fences were installed in two of the primary forest transects from March to April 1999. VES and AES are frequently used for rapid assessments and the evaluation of larger areas. VES and AES thus could be used to evaluate if the randomly chosen transects represented the regional species pool. Trapping with pitfall or funnel traps along drift fences. From February 1999 to December 2000 we compiled 382. during SATS it was often impossible to determine individual parameters. This technique has been formalized as the time-constrained technique of Campbell & Christman (1982). 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. For that reason a maximum recording distance of 12. Coding schemes used for individual recognition were not applied. Thus. The resulting data are expressed in numbers of individuals of a certain species found in an area per unit time. other than sex and species. The width of the acoustic transect depends on the ability to detect each species’ advertisement call. In those cases. This corresponds to 765 transect walks. Dodd 1991). keeping the transect routine identical for both techniques.5 m high and stapled vertically onto wooden stakes. Capture success may vary greatly between species (Corn & Bury 1990. Ptychadena species) are more difficult to trap than terrestrial species that lack these abilities (e. despite their potential abundance. 1997). An array of fences and traps consisted of a central trap (buckets: . data can be expressed in numbers of individuals per time and surface units. In the transect corners. This device is useful to determine species richness of epigaeic organisms (Corn 1994). Anurans that are strong jumpers (e.5 4 hours of transect sampling. For practical reasons “man-hours” can be used. Drift fences consisted of durable green plastic gauze. For both SVTS and SATS. this behavior can be exploited for acoustic monitoring.RÖDEL & ERNST toe clipping (Donnelly et al. At capturing sites a thorough description of the microhabitat was recorded. Visual (VES) and acoustic (AES) encounter surveys.g. An area or habitat is searched systematically for individuals in a defined time period. Standardized acoustic transect sampling (SATS).g. Since in the majority of frog species males use species-specific calls to advertise their position to potential mates and rivals (Wells 1977).
SATS = species recorded during acoustic transect walks. we applied that method during a herpetological survey of the Haute Dodo and Cavally forests. They were checked at least every morning. fossorial and aquatic species independent of families. unpubl. Each segment was tightened around a plastic bucket with an opening angle of 45°. In the course of this study. Sampling success is referred to as number of species within families recorded. data). as sampling success and efficiency may vary with regard to the particular biology of a species. and in a forest fragment (Paulé-Oula 2) outside TNP from 30 August to 12 September 2002 (A. which are situated south and west of TNP respectively (Rödel & Branch 2002. 1. 5 Ta ïF . Hillers et al. using a particular sampling method. Numbers above bars represent total number of species recorded with the respective method. In addition. Number of species per family recorded using different sampling methods. Trap = pitfall and funnel traps. TC = species recorded during transect walks (SVTS and SATS combined). arboreal.. 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). The data from pitfall and funnel traps can be expressed as number of individuals per trap-day. TaïF = all species recorded with all methods in forest habitats around SRET station. results were analyzed for leaf litter. SVTS = species recorded during visual transect walks. 220 mm bottom diameter) and two triangular fence segments (total length 16 m). Traps were checked at least on a daily basis. FIG. The ends of each segment were flanked with additional plastic buckets. AES/VES = visual and acoustic encounter surveys (only species that have been recorded in forest habitats around SRET station). Branch & Rödel 2003). Funnel traps summed to a total of 384 trap-days. Pitfall trapping time summed to a total of approximately 4000 trap-days. one on either side. 285 mm top diameter. Additionally. funnel traps (see Branch & Rödel 2003) were installed in a primary forest transect from 18 to 27 September 2002. Duct tape was used to reduce the diameter of the buckets and construct funnel-like openings in order to impede escaping from traps.AMPHIBIAN MONITORING IN TROPICAL FORESTS 275 mm deep.
. p = 0. Ten species (23. sp. B. df = 5.3 % of TNP species).7 %) of nine families. thus making overall sampling success very high (Tab. P. Hoplobatrachus occipitalis.0 % of TaïF) in eight families. Sampling success and data quality varied between different methods. Effectiveness in detecting species within given families differed significantly between methods (Friedman test. TC. and P. and between different amphibian families or guilds (Figs.g. Numbers above bars represent total number of species recorded with the respective method. RESULTS In total. Bufo superciliaris) or simply absence in the area under investigation (e. Trap). Seven species (Bufo maculatus. SATS) we recorded a total of 15007 individual amphibians belonging to 37 species (86. 43 species were recorded in forest habitats within the study region (Fig. nienokouensis. 1. 2003. SATS. Appendix 1). 2). bibroni.RÖDEL & ERNST FIG. mascareniensis. 2. 1.g. SVTS. Ptychadena pumilio. H.002. 2. Sampling efficiency in families containing higher numbers of genera and/or species was only marginally lower. Rödel et al. χ2 = 19. clearing around SRET) and thus ignored in the analysis.2 %) were recorded with pitfall and funnel trapping. compare Appendix 1). Phlyctimantis boulengeri ). P. 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. Phrynobatrachus fraterculus. Number of species per amphibian guild recorded using different sampling methods. Phrynobatrachus calcaratus.) were recorded exclusively outside forest habitats (e. Taking all methods into account. H. VES/AES revealed 39 forest species (90. two species were each listed in two guilds (compare Appendix). Three leaf litter species (Amnirana occidentalis.) or pitfalls only (Geotrypetes seraphini). N = 10.199. Perret 1988. 2002. 56 amphibian species are known to occur in TNP (Schiøtz 1967..g. annulatus) were exclusively recorded during transect walks. 1. regularis. rather than actual sampling method insufficiency. Rödel & Ernst 2000. . AES/VES. Failure to detect particular species in this region was most probably due to their general scarcity (e. Four arboreal frogs were exclusively recorded with these methods (Hyperolius picturatus. P. fusciventris. Two fossorial species were exclusively detected either with VES and pitfall traps (Hemisus sp. Hyperolius guttulatus). see Appendix 1). tested for TaïF.. For abbreviations see Fig. During transect sampling (SVTS. Rödel 2000.
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).8 0.0 100. Reference is the number of species that were recorded using all methods in forested habitats around the SRET station (TaïF).0 50. Trap).5 100. unpubl.0 0.0 Trap 100.7 % of recorded arboreal species).0 100. data).8 0.0 100.0 100. Diurnal ground-dwelling frogs were best recorded quantitatively using SVTS. asytlosternids) were best recorded with SATS. In TNP funnel traps were ineffective. p = 0. Branch & Rödel 2003).0 33.0 100.0 100.2 66. was difficult as detectability of species varied considerably with (e..0 100. Most arboreal frogs could be recorded with SVTS.0 76.0 100.0 36.0 100.0 100.0 100.0 AES/VES 0. We only captured a few leaf litter frogs. which is mute.0 72.0 100.0 100.0 0. Likewise.8 0.0 100. 7 . As in TNP. 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. With the exception of Acanthixalus sonjae. A distance of 1 m (visual) and 12.5 100.067.0 100. χ2 = 10.5 m (acoustic) proved to work equally well in all habitats investigated. neither between given family or guild. SATS. No additional species were detected with funnel traps.47 specimens per trap-day in contrast to 0.7 100.0 100.0 77.0 0.4 100. 1. In contrast.0 SATS 0. VES/AES. but that concerned mostly single specimens. Quantification of these data. Schaich & Hamerle post hoc multiple comparisons revealed no significant differences between methods at α = 0.7 0.0 68.0 31.7 0. TC.g. the fossorial Geotrypetes seraphini was recorded with pitfall traps only. N = 4.0 100.0 76.0 100. presence and abundance of purely aquatic species and tadpoles of all species were underestimated with all methods used here.0 0.7 100.0 40.294. As aquatic sites were only taken into account. All other species were recorded with VES/AES (Rödel & Branch 2002.3 We could not detect a statistical difference in sampling efficiency between different amphibian guilds using the methods employed (Friedman test.0 0.0 77.0 50. Most specimens were captured with funnel traps (0. Percentage of species within an amphibian family or guild that were recorded using a particular method.05.0 100. df = 4.0 100. other than based on time units. 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 0.0 80.0 64. Trapping success was highest in leaf litter frogs (56. VES and AES revealed highest species numbers.0 100.4 0.0 100.8 100.03 specimens per trap-day in pitfall traps). transect walks made it possible to search for amphibians in a very comparative way. 1990).) vegetation density. tested for TaïF.0 SVTS 0. but very low in arboreal species (6. when being a part of the transect line itself. in a primary forest transect with none in the forest fragment (Hillers et al.0 100.0 0.0 100.0 100.0 66. quantitative data for all arboreal species were best assembled with SATS.0 81.0 0.0 100. SVTS. However.0 100. For other abbreviations see text and Fig. 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 86.5 % of recorded leaf litter frogs). all were recorded using other methods as well.AMPHIBIAN MONITORING IN TROPICAL FORESTS TABLE 2. nocturnal leaf litter frogs (arthloeptids.
Although widely used and recommended. but appear not to be appropriate when standardized quantitative sampling is required. This was also true for most of the arboreal species belonging to the families Hyperoliidae and Rhacophoridae. such as the very abundant Arthroleptis sp. (1995) a simple comparison of species numbers may not be sufficient to evaluate the efficiency of different sampling methods. We found it more realistic to apply a (clearly subjective) measure for habitat complexity on which to base collection effort during VES/AES. The percentage of species within supraspecific taxonomic units.. inaccessible areas than in areas that are easy to monitor. and iv: environmental impact. Methods of collecting quantitative data for aquatic amphibians and for tadpole assemblages have been described by Shaffer et al. Some taxa. When including relative abundance measurements these methods were insufficient. such as species groups.g.RÖDEL & ERNST DISCUSSION Choice of methods should reflect the best compromise between i: practicability. Although the combination of AES and VES provided an adequate approximation to real conditions (see also Doan 2003). ii: specific aims. especially arboreal and fossorial. Cardioglossa leucomystax. pitfall traps and drift fences . 1996). Rödel 2003. which are mute (Drewes 1984. genera or families. The combined transect sampling methods provided a close approximation to the real presence and abundance of species. VES and AES are useful tools for the compilation of species inventory lists.g. during rapid assessment surveys (for recent West African examples see Rödel & Branch 2002.. They suggest using relative species richness within taxonomic entities. these methods do not seem to sample a given fauna adequately well (Allmon 1991. Thus even time-based quantification of these data is questionable. as well as SVTS alone. a combination of SVTS and SATS. In addition. Doan 2003). but is certainly just as well suited for the investigation of tropical anuran assemblages in general. (1997) and Rödel (1998a). in order to sample particularly secretive leaf litter species. Pitfall trapping was only useful in detecting fossorial species that were not encountered during transect walks. We found it impossible to quantify VES and AES other than in relation to time. Standard plot sampling and total removal plots were not tested. Whereas transect walks provide the possibility of monitoring amphibians in a very comparative way. Simple presence/absence data of aquatic species and stages can best be gathered by direct observation (surfacing individuals) and with dip-netting. can then be compared to the real conditions in an area or habitat. acoustic sampling appeared to be indispensable since these species were more readily detected by their calls than 8 by sight. this only holds true if species numbers alone are considered. and will likewise hold true for other arboreal anurans throughout the tropics. According to Pearman et al. were nearly as successful in detecting species. Migrating pipid frogs can also be easily recorded with pitfall and funnel traps. could be detected exclusively using visual sampling methods. data acquisition varies considerably depending on habitat type in VES and even AES.2. 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. Olson et al. Schiøtz 1999. The methods VES and AES in combination with pitfall traps with drift fences were useful in sampling additional. We found no additional species and only very few specimens by funnel trapping. Bury & Corn 1987. 2003). SVTS was generally the best method for detecting diurnal leaf litter frogs. Nonetheless. Rödel & Ernst 2003). In this study this has been done by determining relative species richness with reference to families or guilds. However. However. VES and AES yielded highest species numbers. (1994). Rödel et al. such as species of the genus Acanthixalus. Therefore transect sampling can be considered to be the method of choice when sampling leaf litter anurans. Arthroleptis sp. as an adequate measure. e. and failed to detect aquatic and fossorial species. especially in long-term field studies within temperate regions (e. In general SATS was very efficient in detecting nocturnal leaf litter frogs and treefrogs but less successful in detecting diurnal leaf litter frogs. The differences in efficiency of visual versus acoustic sampling varied between taxa. iii: efficiency. or Phrynobatrachus alticola. The methods tested within this study proved to perform with varying degrees of efficiency. Semlitsch et al. The advantage of transect sampling was especially obvious when focusing on leaf litter species. spending more time in complex. as they impose a severe disturbance on the system under investigation and thus were not suitable to our goals. such as the Hylidae. whereas visual sampling appeared to be superior to acoustic sampling. determined by using a particular sampling method. Vonesh 2001. species not encountered during transect sampling.1.
A comparison of trapping data with data other than those of the observer in question seems to be difficult. If the investigation is interrupted for some time. if not on amphibians then on other organisms. as trapping success per trap per time unit. With the exceptions of aquatic and fossorial species they pro- vided data that cover the whole community very well. Rodda et al.. Traps have to be checked at least daily. none of them uniquely found by this method. experience of investigator and prevailing weather conditions (see Branch & Rödel 2003). generalization of data with regard to phenology or abundances is less possible. 1. Vonesh 2001. However only two species. for other abbreviations see text and Fig.g. however. However. Thus data should be gathered throughout a whole season to obtain a thorough knowledge of the local fauna. e. as well as environmental changes. traps have to be uninstalled or closed and reopened when starting again. VES and AES had probably the lowest environmental impact. Parris 1999. the frequency of sampling can be reduced when general phenological traits of the species recorded have been clarified. Doan 2003). The impact of traps and drift fences was difficult to judge.AMPHIBIAN MONITORING IN TROPICAL FORESTS proved to be the least effective method. Donnelly et al. (2004) had high trapping success with pitfalls in Gabon when considering numbers of individuals. 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 . However trapping success depends very much on choice of site. 2001. one should test the efficiency at a particular site in advance before installing these traps on a broad scale. Data can be used for comparisons between habitats. VES and AES are less time consuming. Burger et al. in theory. SVTS and SATS provided data on species and habitats und thus may reveal changes in composition and species abundance. this method cannot be recommended for standardized sampling of tropical anurans. amphibian guild and environmental impact. Effectiveness of methodology in regard to type of data required.g. Costs with regard to time spent for data acquisition was highest in trapping and in transect walks. They collected only six species (out of a total of 132 species of amphibians and reptiles being recorded).) that leopard (Panthera pardus) tracks TABLE 3. the fossorial Hemisus perreti and the aquatic Silurana epitropicalis. in consecutive field seasons. When considering the relatively high costs and time investment due to the high level of maintenance required (Parris 1999). Data from both trapping methods can. Transects will have an environmental impact.6 % of all individuals captured with this method. In our experience. We observed (e. QS/TR = quadrat sampling/total removal sampling (data from Allmon 1991. Transect walks are time intensive. SVTS and SATS have been proven to be the only methods that provided quantitative data on forest amphibians with regard to space and time. seasons and years. (2001) report on similar experiences using this method in a herpetological survey in Guyana. be quantified in relation to time. Theoretically it might well be that trapping will result in dislocating at least some species from the area where traps have been established. accounted for 72. It is advisable to perform them randomly independent of prevailing weather conditions.
R. Gabon: Species turnover along an elevational gradient. R. 1991. Rev.R. Observations sur la reproduction et la dynamique des populations de quelques anoures tropicaux I. Ecol. However. possibly accompanied by opportunistic trapping.. J. & S. Herpetological survey of the Haute Dodo and Cavally forests. Rödel. & M. Ann. fond. consequent long-term effects on forest structure. L. Standard methods for amphibians. G. Hammond. not only on their breeding sites but throughout the whole range of habitats used by them. Salamandra 39: 91–110. 1978. Phrynobatrachus accraensis. 1994. & S.. Chatelain.J. U.S. Central Amazon.. Project W08 BIOTA-West. 1964. Y. 1982.. W. Channing.” of the Republic of Ivory Coast. Burger. Biodiv. and C. 939 pp. Field techniques for herpetofaunal community analysis.R. Glos.C. Smithsonian Institution Press.P. Afr. M...A. Jr. A plot study of forest floor litter frogs. Barbault. 20: 64–77. & A. Olson et al.. Berlin a. for specific questions (cf..S. 1987. 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. Grundzüge der Vegetationskunde. Memoir 28: 145–186. Canopy fogging of an overstory tree – recommendations for standardization. Trefaut Rodrigues. A 41: 417–428. Pp. Gbamlin. W. western Ivory Coast..D.. Advantages and disadvantages of different methods are summarized in Tab... Global amphibian declines: a problem in applied ecology. Washington & London. N’Dri. Transect cutting and the use of transects should therefore be done with the necessary precautions. Braun-Blanquet. 2003. (eds.M.J.Côte d’Ivoire helped with transportation. R. Hillcoat provided valuable comments on the manuscript.. Richards. Alford.S. Boehnke. Foster. P. Furthermore transects provide an effective method of investigating leaf litter frogs in particular. The access permit to TNP was issued by the “Ministère de la Construction et de l’Environnement. Ouoro provided invaluable help during fieldwork. 1979a. Trefaut Rodrigues. 1998. Research permission was given by the “Ministère de l’Enseignement Supérieur et de la Recherche Scientifique.. Christman.. 1994. 364 pp. Donnelly. Allmon. REFERENCES Adis.G. J. Bortz. SVTS and SATS are especially useful for long-term studies. Barbault. in the worst case. W. Wildlife Management 51: 112–119. TROPENBOS . 7: 503–522. Department of the Interior. Calif. & M. 1997. 2003) transects should be supplemented by specifically investigating aquatic breeding sites with the appropriate set of standardized techniques (Heyer et al. U.1964. & R. ACKNOWLEDGMENTS This study is part of the BIOLOG-program of the German Ministry of Education and Science (BMBF.. Ecol. N. Basset. T. noire. Lienert. 1979b. Trop.): Herpetological communities. All this support is gratefully acknowledged.-O. Trop. Corn. Barbault. A.. Wien. (ed.): Measuring and monitoring biological diversity. Fish and Wildlife Service 13. and B. 1999. Ecotropica 4: 93–97.W.. Ptychadena maccarthyensis et Ptychadena oxyrhynchus. Rödel 1998a). Hayek.-A.RÖDEL & ERNST were more often seen on the transect path than in undisturbed forest. L. W.o. Spichiger. 109–117 in Heyer. Arthroleptis poecilonotus. Pflanzensoziologie. Bull.” G. 2 ed. Part II: Trapping results and reptiles.A. Hofer. Ecol. Branch. & M. H. 3. & K. & M. 01 LC0017). Evaluation of pitfall trapping in northwestern forests: trap arrays with drift fences..B. J. Pp. Trefaut Rodrigues.. Amphibians and reptiles of Monts Doudou. 5: 37–53. Conserv. Floren. J. For simple short-term surveys we recommend VES and AES. Observations sur la reproduction et la dynamique des populations de quelques anoures tropicaux IV. J. Gautier. Wildlife Research Report 13. Syst. 1990. Inst.. P. 2004.A. Observations sur la reproduction et la dynamique des populations de quelques anoures tropicaux III. thus generating a much more complete picture of an amphibian community than is possible with other methods. 30: 133–165. Brazil. 1996. Bury. and the “Taï Monkey Project” provided logistic support..Y. Terre Vie 32: 441–452. R. & M. Halliday. The “Projet Autonome pour la Conservation du Parc National de Taï”. A recent history of forest fragmentation in southwestern Ivory Coast.” Working in TNP was kindly permitted by the Commandant K. Sci. Campbell. 193–200 in Scott. a well visible path becomes established that at least in primary forests remains so for years. After using transects intensively for some time. C. 865 pp. Linsenmair. 10 .E. & K. R.. This might affect densities of other mammals including possible seed dispersers with. The presented transect design proved to be adequate for representative sampling and appeared to be appropriate for most studies involving multivariate structure data. M.R. Branch.S. McDiarmid. Corn. R. Acad. Lips et al. Springer Verlag. Verteilungsfreie Methoden der Biostatistik. Sér.W. J. & P. Straight-line drift fences and pitfall traps. Springer Verlag.
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7°10’ W). l = leaf litter species. 1 = known from TNP (Schiøtz 1967. 3 = Phrynobatrachus guineensis is a leaf litter frog but reproduces in small water-filled tree holes (Rödel 1998b). VES/AES = species recorded via visual and acoustic encounter surveys in forest habitats near SRET. 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. SRET = species recorded with VES/AES on the clearing around the SRET station. Perret 1988) but not recorded by us. 5°25’ N. 6 = recorded north of area under investigation. 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. 2 = determination not possible at present time. G = species recorded in southern TNP (Guiroutou. TP = all species recorded on transects in primary forest. List of amphibian species recorded in Taï National Park (TNP). TaïF = species recorded in forest habitats around the SRET station. aq = aquatic species. TC = all species recorded on transects. f = fossorial species.AMPHIBIAN MONITORING IN TROPICAL FORESTS APPENDIX. a = arboreal species. 4 = Acanthixalus sonjae lives aquatically in large water filled tree holes but forages in surrounding vegetation (Rödel et al. TS = all species recorded on transects in secondary forest. 2003). species Gymnophiona Caecilidae Geotrypetes seraphini Anura Pipidae Silurana tropicalis Bufonidae Bufo regularis Bufo maculatus Bufo togoensis Bufo taiensis Bufo superciliaris Hemisotidae Hemisus sp. 5 = treated as full species and not as subspecies of Hyperolius fusciventris due to sympatric occurrence in TNP (see Rödel & Branch 2002). a l l l l l 13 .
12 Arthroleptis sp. aq a 14 .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.
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