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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
Transects were arranged in pairs. 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. Vonesh 2001. Formenty pers. A number of recently published regional field monitoring manuals for amphibian surveys (Latin America: Lips et al. b. that stretches west of the Dahomey Gap from Ghana into Sierra Leone and Guinea (Schiøtz 1967. consists of a series of small squares (quadrats) that are laid out at randomly selected sites within a habitat then thoroughly searched for amphibians. Rodda et al. The major rainy season stretches from September to October. Riezebos et al. Given the lack of time for the development of effective conservation programs. data: Taï Monkey Project).RÖDEL & ERNST essential for academic and practical purposes alike. A minor rainy season lasts from March/April to July. standardization of methods that yield broad-scale comparative data is now needed more than ever. Floristically the TNP belongs to the Guinean-Congo-Region (Guillaumet 1967. 2001) were a priori excluded from testing. Parren & de Graaf 1995. 5°50’N. 7°20’W). and iv) has a low environmental impact. The starting coordinate for each transect always marked the southeast corner to ensure identical geographic orientation between sampling units. such as standard plot sampling (Allmon 1991. We thereby evaluated several methods that are already commonly used and present a set of techniques that. The second. (1994) and PACPNT (2000). Doan 2003) or total removal plots (Barbault & Trefaut Rodrigues 1978. Every subunit was marked with numbered colored flag-tape. Precipitation is distributed across two more or less distinct periods. TNP is the largest remaining protected area of rain forest in West Africa. followed by a short dry season in July/August.3 km. proved to be most effective for forest amphibians. The maximum distance between transects was 6. Transect paths were kept open so that walking at a constant speed was possible at all times. 1979a. Each had a north-south extension of 200 m and an east-west extension of 100 m. situated in western Côte d’Ivoire (5°08’6°07’N. Thus widely used techniques. Richards 1996). The first. as they may result in severe disturbance to the system under investigation. 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. For data acquisition the complete transect length of 600 m was subdivided in 25 m subunits (24 subunits/transect). 2001. known as quadrat sampling (plot design). also for para-ecologists. Côte d’Ivoire. 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. Mean annual precipitation in the study area was 1806 mm during 1988–2002 (± 297mm. 1994. uses linear transects instead of squares. ratures vary between 20 and 33° C. Lawson 1986. Parts of the habitats on the park’s periphery were subject to more or less intense logging and cultivation until 1998 (P. The core dry season lasts from November until February/March. More detailed descriptions of TNP are provided by Rompaey (1993). map “Projet OMS Forêt de Taï”). thus ensuring that all habitat types of a certain area within the very inhomogeneous forest were covered. four in secondary forest). Whereas quadrat sampling can be used to determine the species present in a homogenous area. 6°47’-7°25’W). The minimum distance between adjacent transects was 200 m. as well as their relative abundances and densities. Africa: Howell 2002) reflect this need. and unpubl. PACPNT 2000). We avoided extensive cutting and thus manipulation of important habitat features. iii) is efficient concerning time and money. The rectangular transect design is a combination of two widely used standard techniques. All evaluations and recommendations are based on a study of tropical amphibian assemblages conducted over four years (1999–2002) in Taï National Park. known as transect sampling. transect sampling can provide similar data across habitat or dis- MATERIAL AND METHODS Study site The study was conducted in Taï National Park (TNP). Poynton 1999). Daily tempe2 . that ii) is easy to handle. comm. It can be characterized as humid tropic seasonal (Riezebos et al. The mean annual temperature is about 25° C (Rompaey 1993). Our study site was located 23 km southeast of the small town of Taï and comprised about 30 km2 of primary and secondary rain forest around the Station de Recherche en Ecologie de Taï (SRET. on the basis of our experience in the field as well as with consecutive data analyses. 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.
35 m/s). To assess habitat preferences of particular species (e. surface and depth. Pearman 1997).30–0. lower tree stratum: 3–10 m. understory: < 0. We assumed that the number of plants with small dbh is greater in degraded. rectangular ones provide the possibility of easily up. Specimens heard but not seen within this distance were not searched for and not included in the SVTS data sets (see below). Categories of habitat variables measured on transects. bush and shrub stratum: 0.or downscaling data without neglecting local habitat diversity (e. definition category 1 1.5 2 2. vegetation density was measured in four strata (see text).g.AMPHIBIAN MONITORING IN TROPICAL FORESTS turbance gradients (Jaeger 1994. thus providing data that are useful to test for sex. Transects were intensively patrolled at a constant speed (0. 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. to assign them to particular guilds).5 3 3.5 m.. both in time and space.5 m). thereby recording all amphibians within a distance of 100 cm from either side of the path. substrate moisture was determined in four categories during every transect walk. 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. and to investigate correlations of species assemblages with environmental variables. 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. Sampling was interrupted for the duration of the recording in order not to overestimate locations in which animals had previously been captured. Sampling methods and sampling effort Habitat characterization. we characterized all habitats using several variables that were recorded at two defined points within each 25 m subunit (beginning and midpoint). In order to quantify the availability of potential aquatic breeding sites. Standardized visual transect sampling (SVTS). divided into seven categories corresponding to particular densities. Compared to linear transects. Definitions of habitat variables are summarized in Table 1. whereas primary for- ests show increasing numbers of plants of larger dbh (Chatelain et al. Edaphic parameters were registered as general substrate types according to Lieberoth (1982).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 . In addition.or age-specific differences in habitat use.5–1. dbh = diameter of plants at breast height. A maximum of four transects was patrolled at night. Jaeger & Inger 1994).g. Usually four to six transects per day were sampled during daytime. Repeated controls of identical transects on consecutive days were avoided to ensure independence of samples.. 1996. Additionally. captured frogs were marked by TABLE 1. for comparison of local with regional species pools). To avoid duplicate recordings. 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. secondary forests. dbh). Sampling was performed independent of prevailing weather conditions. As far as possible all individuals were captured. For a discussion of the advantages of rectangular sample units over quadratic or circular ones see Krebs (1989) and McCune & Grace (2002).
The resulting data are expressed in numbers of individuals of a certain species found in an area per unit time. keeping the transect routine identical for both techniques.g. 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). Recaptures were excluded from the analyses.. this technique allows the detecting of cryptic species that. Furthermore.and AES-effort was kept comparable to the time spent on SVTS and SATS surveys throughout the entire study period. and to estimate relative abundances of species within an assemblage (Crump & Scott 1994). 1997). It can be used to determine the species richness of an area and the species composition of a local assemblage.. 0. From February 1999 to December 2000 we compiled 382. Due to the simplicity of the method. Ptychadena species) are more difficult to trap than terrestrial species that lack these abilities (e. In the transect corners. VES can only provide information on the presence or absence of a species in an area but are inadequate for determining abundances. other than sex and species.e. Coding schemes used for individual recognition were not applied. For practical reasons “man-hours” can be used. despite their potential abundance. Thus. At capturing sites a thorough description of the microhabitat was recorded. individual marking should be restricted to studies in which individual recognition is indispensable. VES and AES are frequently used for rapid assessments and the evaluation of larger areas. Trapping with pitfall or funnel traps along drift fences. An array of fences and traps consisted of a central trap (buckets: . This corresponds to 765 transect walks. beginning with segment 1: segment 1. and as the time-constrained searches of Corn & Bury (1990). Counts can be used to estimate relative abundance of calling males. during SATS it was often impossible to determine individual parameters. Pitfall traps with drift fences were installed in two of the primary forest transects from March to April 1999. Likewise microhabitat description was not always possible.5 m to either side of the transect was defined. 1994. Total VES. For that reason a maximum recording distance of 12. data can be expressed in numbers of individuals per time and surface units. 12 and 20). VES and AES thus could be used to evaluate if the randomly chosen transects represented the regional species pool. 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. Standardized acoustic transect sampling (SATS). and additionally in only one of these transects from January 2001 to September 2002. which can be adjusted to the complexity of the habitats being sampled. We used these methods continuously in all habitats of the study area including the transect areas. As opposed to visual sampling. as well as breeding habitat use and breeding phenology of species (Zimmerman 1994). For both SVTS and SATS. Drift fences consisted of durable green plastic gauze. following the characterization routine used for general habitat description.. The width of the acoustic transect depends on the ability to detect each species’ advertisement call. 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). Since in the majority of frog species males use species-specific calls to advertise their position to potential mates and rivals (Wells 1977). this behavior can be exploited for acoustic monitoring.g. According to Corn & Bury (1990). Anurans that are strong jumpers (e. In those cases.5 m high and stapled vertically onto wooden stakes.RÖDEL & ERNST toe clipping (Donnelly et al. species composition. habitat analyses can be based on data obtained from the general habitat characterization for single transects or segments. Throughout transect walks a combination of visual and acoustic techniques was applied at all times. Capture success may vary greatly between species (Corn & Bury 1990. Visual (VES) and acoustic (AES) encounter surveys. This device is useful to determine species richness of epigaeic organisms (Corn 1994). Bufo species). Individuals below nine mm SVL were not marked due to their small size. Dodd 1991). In concordance with these authors the methods were only used as a qualitative or semi-quantitative tool within the scope of our studies. An area or habitat is searched systematically for individuals in a defined time period. Henle et al. Calls from greater distances cannot be unambiguously identified and the chance of duplicate recordings in neighboring segments increases.5 4 hours of transect sampling. 8. thus creating 25 m x 25 m acoustic sampling plots. may be underestimated when exclusively using visual techniques. This technique has been formalized as the time-constrained technique of Campbell & Christman (1982).
Additionally. Sampling success is referred to as number of species within families recorded. funnel traps (see Branch & Rödel 2003) were installed in a primary forest transect from 18 to 27 September 2002. In the course of this study. SATS = species recorded during acoustic transect walks. arboreal. TaïF = all species recorded with all methods in forest habitats around SRET station. Data from each sampling method were compared with each other and to a list of species records that we gathered through January 1999 to September 2002 for the forest parts of the SRET region (Appendix 1). Number of species per family recorded using different sampling methods. data). 285 mm top diameter. In addition. using a particular sampling method. Pitfall trapping time summed to a total of approximately 4000 trap-days. AES/VES = visual and acoustic encounter surveys (only species that have been recorded in forest habitats around SRET station). unpubl. 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. Duct tape was used to reduce the diameter of the buckets and construct funnel-like openings in order to impede escaping from traps. Trap = pitfall and funnel traps. Traps were checked at least on a daily basis. 220 mm bottom diameter) and two triangular fence segments (total length 16 m). The data from pitfall and funnel traps can be expressed as number of individuals per trap-day. one on either side. SVTS = species recorded during visual transect walks. we applied that method during a herpetological survey of the Haute Dodo and Cavally forests. TC = species recorded during transect walks (SVTS and SATS combined). Branch & Rödel 2003). They were checked at least every morning. 1. Hillers et al. results were analyzed for leaf litter.AMPHIBIAN MONITORING IN TROPICAL FORESTS 275 mm deep. Funnel traps summed to a total of 384 trap-days.. fossorial and aquatic species independent of families. Numbers above bars represent total number of species recorded with the respective method. 5 Ta ïF . which are situated south and west of TNP respectively (Rödel & Branch 2002. FIG. as sampling success and efficiency may vary with regard to the particular biology of a species. The ends of each segment were flanked with additional plastic buckets.
. Number of species per amphibian guild recorded using different sampling methods. SVTS. thus making overall sampling success very high (Tab. p = 0. Seven species (Bufo maculatus. 1. Hoplobatrachus occipitalis. During transect sampling (SVTS. clearing around SRET) and thus ignored in the analysis. H. AES/VES. Three leaf litter species (Amnirana occidentalis.) or pitfalls only (Geotrypetes seraphini). Phrynobatrachus fraterculus. Bufo superciliaris) or simply absence in the area under investigation (e. two species were each listed in two guilds (compare Appendix). regularis. and between different amphibian families or guilds (Figs. Numbers above bars represent total number of species recorded with the respective method. Four arboreal frogs were exclusively recorded with these methods (Hyperolius picturatus.2 %) were recorded with pitfall and funnel trapping. Phlyctimantis boulengeri ).7 %) of nine families. compare Appendix 1). P. N = 10. Taking all methods into account. Rödel et al. χ2 = 19. Ptychadena pumilio.3 % of TNP species).RÖDEL & ERNST FIG. rather than actual sampling method insufficiency. SATS. Two fossorial species were exclusively detected either with VES and pitfall traps (Hemisus sp. 2003. Appendix 1). annulatus) were exclusively recorded during transect walks. RESULTS In total. . mascareniensis. Phrynobatrachus calcaratus. Rödel & Ernst 2000. Hyperolius guttulatus). Failure to detect particular species in this region was most probably due to their general scarcity (e. VES/AES revealed 39 forest species (90.) were recorded exclusively outside forest habitats (e. and P.. 2002. 2. Sampling success and data quality varied between different methods. 43 species were recorded in forest habitats within the study region (Fig. H. sp. Ten species (23. SATS) we recorded a total of 15007 individual amphibians belonging to 37 species (86. B. 56 amphibian species are known to occur in TNP (Schiøtz 1967. 2. TC. tested for TaïF. For abbreviations see Fig.g. 2). 1.199. 1. Perret 1988.g.0 % of TaïF) in eight families. Using all methods we recorded 50 species in the region of the SRET (89. nienokouensis.002. Families containing only 6 a few species in the TNP area showed the highest recording rates. Rödel 2000. fusciventris. bibroni.g. Trap). Effectiveness in detecting species within given families differed significantly between methods (Friedman test. Sampling efficiency in families containing higher numbers of genera and/or species was only marginally lower. see Appendix 1). P. df = 5.. P.
the fossorial Geotrypetes seraphini was recorded with pitfall traps only.0 SATS 0. N = 4.AMPHIBIAN MONITORING IN TROPICAL FORESTS TABLE 2.2 66.0 100. transect walks made it possible to search for amphibians in a very comparative way.0 76.0 100.0 100.0 100. Schaich & Hamerle post hoc multiple comparisons revealed no significant differences between methods at α = 0.0 80. unpubl.5 100. As aquatic sites were only taken into account. As in TNP.0 77. but very low in arboreal species (6.0 SVTS 0.0 100.0 100. However.5 % of recorded leaf litter frogs).0 66.5 m (acoustic) proved to work equally well in all habitats investigated.7 0.0 0. In Haute Dodo and Cavally forests we captured 38 % of the recorded amphibian species (16 of 42) with a combination of pitfall and funnel traps.8 0. Reference is the number of species that were recorded using all methods in forested habitats around the SRET station (TaïF). Trap).05. χ2 = 10. neither between given family or guild.0 33.0 100. No additional species were detected with funnel traps. p = 0.0 100.0 100. all were recorded using other methods as well. For other abbreviations see text and Fig.0 31.0 100. but that concerned mostly single specimens.4 0.0 100. Most arboreal frogs could be recorded with SVTS.067. tested for TaïF.4 100.0 100. nocturnal leaf litter frogs (arthloeptids. Most specimens were captured with funnel traps (0. quantitative data for all arboreal species were best assembled with SATS.0 77.0 76.8 0.0 100.0 100.7 0. SVTS.0 72. 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. df = 4.) vegetation density.8 100. A distance of 1 m (visual) and 12.0 100. 1990). Diurnal ground-dwelling frogs were best recorded quantitatively using SVTS.0 50. 1. TC. Branch & Rödel 2003).0 100.294. 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. VES/AES.0 0.0 100.0 36.5 100. asytlosternids) were best recorded with SATS. Quantification of these data.0 100.0 AES/VES 0.g. in a primary forest transect with none in the forest fragment (Hillers et al.0 100.7 100.0 0.0 100.0 0. other than based on time units. In contrast.0 100.03 specimens per trap-day in pitfall traps). when being a part of the transect line itself.0 100.0 100. With the exception of Acanthixalus sonjae.0 0.0 40. All other species were recorded with VES/AES (Rödel & Branch 2002. Percentage of species within an amphibian family or guild that were recorded using a particular method. VES and AES revealed highest species numbers. SATS. presence and abundance of purely aquatic species and tadpoles of all species were underestimated with all methods used here.0 64. We only captured a few leaf litter frogs.0 Trap 100. data).0 100.8 0. Trapping success was highest in leaf litter frogs (56. Likewise. which is mute.0 81.0 0.3 We could not detect a statistical difference in sampling efficiency between different amphibian guilds using the methods employed (Friedman test.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).0 100.0 86.0 0.0 68.0 0.7 100.0 50. In TNP funnel traps were ineffective.7 % of recorded arboreal species). 7 ..47 specimens per trap-day in contrast to 0. was difficult as detectability of species varied considerably with (e.
We found it impossible to quantify VES and AES other than in relation to time. However. Standard plot sampling and total removal plots were not tested. whereas visual sampling appeared to be superior to acoustic sampling. 2003). Therefore transect sampling can be considered to be the method of choice when sampling leaf litter anurans. The advantage of transect sampling was especially obvious when focusing on leaf litter species.. This was also true for most of the arboreal species belonging to the families Hyperoliidae and Rhacophoridae. which are mute (Drewes 1984. can then be compared to the real conditions in an area or habitat. The methods VES and AES in combination with pitfall traps with drift fences were useful in sampling additional. They suggest using relative species richness within taxonomic entities. Semlitsch et al. Olson et al. inaccessible areas than in areas that are easy to monitor. and iv: environmental impact. e. Bury & Corn 1987. Rödel & Ernst 2003). The combined transect sampling methods provided a close approximation to the real presence and abundance of species. In this study this has been done by determining relative species richness with reference to families or guilds. Whereas transect walks provide the possibility of monitoring amphibians in a very comparative way. Migrating pipid frogs can also be easily recorded with pitfall and funnel traps. Pitfall trapping was only useful in detecting fossorial species that were not encountered during transect walks. Methods of collecting quantitative data for aquatic amphibians and for tadpole assemblages have been described by Shaffer et al. were nearly as successful in detecting species. such as the Hylidae. When including relative abundance measurements these methods were insufficient. species not encountered during transect sampling.. According to Pearman et al. in order to sample particularly secretive leaf litter species. or Phrynobatrachus alticola. such as species of the genus Acanthixalus. 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. as they impose a severe disturbance on the system under investigation and thus were not suitable to our goals. such as species groups. during rapid assessment surveys (for recent West African examples see Rödel & Branch 2002. Cardioglossa leucomystax. but appear not to be appropriate when standardized quantitative sampling is required. especially in long-term field studies within temperate regions (e. Although the combination of AES and VES provided an adequate approximation to real conditions (see also Doan 2003). especially arboreal and fossorial.2. (1995) a simple comparison of species numbers may not be sufficient to evaluate the efficiency of different sampling methods. these methods do not seem to sample a given fauna adequately well (Allmon 1991. iii: efficiency. but is certainly just as well suited for the investigation of tropical anuran assemblages in general. The differences in efficiency of visual versus acoustic sampling varied between taxa. Nonetheless. Vonesh 2001. Some taxa. determined by using a particular sampling method. The methods tested within this study proved to perform with varying degrees of efficiency.1. a combination of SVTS and SATS. this only holds true if species numbers alone are considered. In general SATS was very efficient in detecting nocturnal leaf litter frogs and treefrogs but less successful in detecting diurnal leaf litter frogs. Schiøtz 1999.g. such as the very abundant Arthroleptis sp. We found no additional species and only very few specimens by funnel trapping. genera or families.RÖDEL & ERNST DISCUSSION Choice of methods should reflect the best compromise between i: practicability. acoustic sampling appeared to be indispensable since these species were more readily detected by their calls than 8 by sight. ii: specific aims. and will likewise hold true for other arboreal anurans throughout the tropics. data acquisition varies considerably depending on habitat type in VES and even AES. The percentage of species within supraspecific taxonomic units. Arthroleptis sp. (1994). and failed to detect aquatic and fossorial species. could be detected exclusively using visual sampling methods. as an adequate measure. VES and AES yielded highest species numbers. (1997) and Rödel (1998a). pitfall traps and drift fences . SVTS was generally the best method for detecting diurnal leaf litter frogs. 1996). Although widely used and recommended.g. Simple presence/absence data of aquatic species and stages can best be gathered by direct observation (surfacing individuals) and with dip-netting. spending more time in complex. Doan 2003). However. as well as SVTS alone. Rödel 2003. In addition. Rödel et al. Thus even time-based quantification of these data is questionable. We found it more realistic to apply a (clearly subjective) measure for habitat complexity on which to base collection effort during VES/AES. VES and AES are useful tools for the compilation of species inventory lists.
We observed (e. A comparison of trapping data with data other than those of the observer in question seems to be difficult. e. In our experience. Theoretically it might well be that trapping will result in dislocating at least some species from the area where traps have been established. if not on amphibians then on other organisms. be quantified in relation to time. none of them uniquely found by this method. VES and AES are less time consuming. one should test the efficiency at a particular site in advance before installing these traps on a broad scale. Vonesh 2001. amphibian guild and environmental impact. Burger et al. the frequency of sampling can be reduced when general phenological traits of the species recorded have been clarified. It is advisable to perform them randomly independent of prevailing weather conditions. for other abbreviations see text and Fig. Traps have to be checked at least daily. If the investigation is interrupted for some time. as trapping success per trap per time unit. the fossorial Hemisus perreti and the aquatic Silurana epitropicalis.. as well as environmental changes. this method cannot be recommended for standardized sampling of tropical anurans. traps have to be uninstalled or closed and reopened when starting again. SVTS and SATS provided data on species and habitats und thus may reveal changes in composition and species abundance. (2004) had high trapping success with pitfalls in Gabon when considering numbers of individuals. Data can be used for comparisons between habitats. (2001) report on similar experiences using this method in a herpetological survey in Guyana. Doan 2003).AMPHIBIAN MONITORING IN TROPICAL FORESTS proved to be the least effective method. SVTS and SATS have been proven to be the only methods that provided quantitative data on forest amphibians with regard to space and time. experience of investigator and prevailing weather conditions (see Branch & Rödel 2003). 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. accounted for 72. Rodda et al.) that leopard (Panthera pardus) tracks TABLE 3. Transect walks are time intensive. 2001. however. They collected only six species (out of a total of 132 species of amphibians and reptiles being recorded). Transects will have an environmental impact. generalization of data with regard to phenology or abundances is less possible. When considering the relatively high costs and time investment due to the high level of maintenance required (Parris 1999).6 % of all individuals captured with this method. The impact of traps and drift fences was difficult to judge. in consecutive field seasons. Donnelly et al. Thus data should be gathered throughout a whole season to obtain a thorough knowledge of the local fauna. Effectiveness of methodology in regard to type of data required. However trapping success depends very much on choice of site. Costs with regard to time spent for data acquisition was highest in trapping and in transect walks. However only two species. 1.g. QS/TR = quadrat sampling/total removal sampling (data from Allmon 1991. in theory. Data from both trapping methods can. Parris 1999. seasons and years.g. VES and AES had probably the lowest environmental impact. With the exceptions of aquatic and fossorial species they pro- vided data that cover the whole community very well.
): Measuring and monitoring biological diversity. Halliday. H. Gbamlin. Central Amazon...” G. Sér. Trefaut Rodrigues. Floren. & A. Lips et al.R.. J. Observations sur la reproduction et la dynamique des populations de quelques anoures tropicaux I.. Verteilungsfreie Methoden der Biostatistik. & S.): Herpetological communities. Barbault.M.o. Olson et al. Ecol. 2003) transects should be supplemented by specifically investigating aquatic breeding sites with the appropriate set of standardized techniques (Heyer et al. C. & R. REFERENCES Adis. ACKNOWLEDGMENTS This study is part of the BIOLOG-program of the German Ministry of Education and Science (BMBF. Inst.S.-O. Linsenmair. Observations sur la reproduction et la dynamique des populations de quelques anoures tropicaux III.” of the Republic of Ivory Coast. Alford.. 1996. The access permit to TNP was issued by the “Ministère de la Construction et de l’Environnement. Barbault. & M. & M. 1997. 10 . 1964. All this support is gratefully acknowledged. R. Trefaut Rodrigues. Global amphibian declines: a problem in applied ecology. & P. W. 2003. Gautier. Barbault.A. 193–200 in Scott. R. Smithsonian Institution Press. and B. Herpetological survey of the Haute Dodo and Cavally forests. Trop. & M. & K. & M.R.. Advantages and disadvantages of different methods are summarized in Tab. Braun-Blanquet. 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. Part II: Trapping results and reptiles. & S. 1994. Branch.1964. Rödel.. Berlin a. R.. 1979b. A recent history of forest fragmentation in southwestern Ivory Coast. 20: 64–77. Basset. Arthroleptis poecilonotus.A.. 1978. Brazil... 3. R. Springer Verlag.J. 1998.. Evaluation of pitfall trapping in northwestern forests: trap arrays with drift fences.W.S. Standard methods for amphibians. M. Amphibians and reptiles of Monts Doudou. Wildlife Research Report 13. consequent long-term effects on forest structure. 2004. and C. Grundzüge der Vegetationskunde. 109–117 in Heyer. Ouoro provided invaluable help during fieldwork. For simple short-term surveys we recommend VES and AES. Canopy fogging of an overstory tree – recommendations for standardization. Afr. McDiarmid. Boehnke. Straight-line drift fences and pitfall traps. (eds. Department of the Interior.P. Transect cutting and the use of transects should therefore be done with the necessary precautions. western Ivory Coast. U. Rödel 1998a). Ptychadena maccarthyensis et Ptychadena oxyrhynchus..R. 5: 37–53.-A. Hayek. A plot study of forest floor litter frogs.C.W. T. noire. Memoir 28: 145–186. Foster. J. J. J. Furthermore transects provide an effective method of investigating leaf litter frogs in particular. Ecol. L. The “Projet Autonome pour la Conservation du Parc National de Taï”. Ecol.Y. 1987. 1979a. 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. 7: 503–522. Sci. Bortz. Salamandra 39: 91–110. Phrynobatrachus accraensis. Corn. J. Lienert. for specific questions (cf.. N’Dri. 01 LC0017). SVTS and SATS are especially useful for long-term studies.S. Chatelain. (ed. Glos. A 41: 417–428. & M.. 1994. W. L. Ann. Channing.G. Hofer. R.. Trefaut Rodrigues. possibly accompanied by opportunistic trapping. in the worst case. N. W. Observations sur la reproduction et la dynamique des populations de quelques anoures tropicaux IV. R. a well visible path becomes established that at least in primary forests remains so for years. Ecotropica 4: 93–97. Conserv. Wien. Hillcoat provided valuable comments on the manuscript.. M. A. Branch. Corn. J. Campbell. P. 1982. not only on their breeding sites but throughout the whole range of habitats used by them.J. TROPENBOS . Gabon: Species turnover along an elevational gradient. Rev. 2 ed.D.. Biodiv. This might affect densities of other mammals including possible seed dispersers with. 364 pp.. thus generating a much more complete picture of an amphibian community than is possible with other methods. Spichiger. Wildlife Management 51: 112–119. 865 pp. 1999. Syst. Jr. Donnelly. & K.. Bull. P. fond.RÖDEL & ERNST were more often seen on the transect path than in undisturbed forest. Project W08 BIOTA-West. After using transects intensively for some time. Hammond. However. Richards. 939 pp. G. and the “Taï Monkey Project” provided logistic support. Pflanzensoziologie. Pp. Terre Vie 32: 441–452.Côte d’Ivoire helped with transportation. Field techniques for herpetofaunal community analysis. Research permission was given by the “Ministère de l’Enseignement Supérieur et de la Recherche Scientifique. Acad.S. Pp. Bury. 1991. Trop.A. Calif.. U. 1990..E.” Working in TNP was kindly permitted by the Commandant K. 30: 133–165.B. Y. Allmon. Christman. Springer Verlag. W. Fish and Wildlife Service 13. Burger.
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TS = all species recorded on transects in secondary forest. l = leaf litter species. a = arboreal species. TP = all species recorded on transects in primary forest. a l l l l l 13 . other than transects. 4 = Acanthixalus sonjae lives aquatically in large water filled tree holes but forages in surrounding vegetation (Rödel et al. Trap = species recorded with pitfall and funnel traps. species Gymnophiona Caecilidae Geotrypetes seraphini Anura Pipidae Silurana tropicalis Bufonidae Bufo regularis Bufo maculatus Bufo togoensis Bufo taiensis Bufo superciliaris Hemisotidae Hemisus sp. 5 = treated as full species and not as subspecies of Hyperolius fusciventris due to sympatric occurrence in TNP (see Rödel & Branch 2002). 6 = recorded north of area under investigation. 7°10’ W). 2 = determination not possible at present time. 2003). f = fossorial species. aq = aquatic species. 5°25’ N. VES/AES = species recorded via visual and acoustic encounter surveys in forest habitats near SRET. G = species recorded in southern TNP (Guiroutou. 1 = known from TNP (Schiøtz 1967. 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. Perret 1988) but not recorded by us. SRET = species recorded with VES/AES on the clearing around the SRET station. 3 = Phrynobatrachus guineensis is a leaf litter frog but reproduces in small water-filled tree holes (Rödel 1998b). 2 Ranidae Hoplobatrachus occipitalis Amnirana albolabris Amnirana occidentalis Aubria occidentalis Ptychadena pumilio Ptychadena bibroni Ptychadena mascareniensis Ptychadena superciliaris Ptychadena aequiplicata Ptychadena longirostris Ptychadena sp. List of amphibian species recorded in Taï National Park (TNP). TC = all species recorded on transects. TaïF = species recorded in forest habitats around the SRET station.AMPHIBIAN MONITORING IN TROPICAL FORESTS APPENDIX.
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.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. aq a 14 .
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