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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
Formenty pers. TNP is part of the Upper Guinea forest block. 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. The mean annual temperature is about 25° C (Rompaey 1993). Lawson 1986. that stretches west of the Dahomey Gap from Ghana into Sierra Leone and Guinea (Schiøtz 1967. The first. known as transect sampling. 2001) were a priori excluded from testing. 1994. known as quadrat sampling (plot design). The maximum distance between transects was 6. Our study sites are either pristine forest areas or have not been logged or cultivated since 1978. 1979a. as they may result in severe disturbance to the system under investigation. 5°50’N. A number of recently published regional field monitoring manuals for amphibian surveys (Latin America: Lips et al. four in secondary forest). situated in western Côte d’Ivoire (5°08’6°07’N. 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. PACPNT 2000). Poynton 1999). Vonesh 2001. Floristically the TNP belongs to the Guinean-Congo-Region (Guillaumet 1967. Precipitation is distributed across two more or less distinct periods. comm. 2001. We aim to present a methodology that i) provides quantitative and qualitative data useful for rapid surveys as well as long-term monitoring programs. Whereas quadrat sampling can be used to determine the species present in a homogenous area. The major rainy season stretches from September to October. Thus widely used techniques.3 km. standardization of methods that yield broad-scale comparative data is now needed more than ever. TNP is the largest remaining protected area of rain forest in West Africa. that ii) is easy to handle. consists of a series of small squares (quadrats) that are laid out at randomly selected sites within a habitat then thoroughly searched for amphibians. Richards 1996). transect sampling can provide similar data across habitat or dis- MATERIAL AND METHODS Study site The study was conducted in Taï National Park (TNP). 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. ratures vary between 20 and 33° C. (1994) and PACPNT (2000). Each had a north-south extension of 200 m and an east-west extension of 100 m. and iv) has a low environmental impact. also for para-ecologists. Riezebos et al. The minimum distance between adjacent transects was 200 m. Every subunit was marked with numbered colored flag-tape. Doan 2003) or total removal plots (Barbault & Trefaut Rodrigues 1978. The starting coordinate for each transect always marked the southeast corner to ensure identical geographic orientation between sampling units. Côte d’Ivoire. Africa: Howell 2002) reflect this need. The second. and unpubl. We thereby evaluated several methods that are already commonly used and present a set of techniques that. Transect paths were kept open so that walking at a constant speed was possible at all times. The core dry season lasts from November until February/March. Rodda et al. b. data: Taï Monkey Project). thus ensuring that all habitat types of a certain area within the very inhomogeneous forest were covered. 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.RÖDEL & ERNST essential for academic and practical purposes alike. Parts of the habitats on the park’s periphery were subject to more or less intense logging and cultivation until 1998 (P. Mean annual precipitation in the study area was 1806 mm during 1988–2002 (± 297mm. on the basis of our experience in the field as well as with consecutive data analyses. A minor rainy season lasts from March/April to July. followed by a short dry season in July/August. We avoided extensive cutting and thus manipulation of important habitat features. It can be characterized as humid tropic seasonal (Riezebos et al. such as standard plot sampling (Allmon 1991. as well as their relative abundances and densities. 6°47’-7°25’W). uses linear transects instead of squares. More detailed descriptions of TNP are provided by Rompaey (1993). 7°20’W). map “Projet OMS Forêt de Taï”). Given the lack of time for the development of effective conservation programs. proved to be most effective for forest amphibians. Daily tempe2 . Parren & de Graaf 1995. Transects were arranged in pairs. For data acquisition the complete transect length of 600 m was subdivided in 25 m subunits (24 subunits/transect). iii) is efficient concerning time and money. 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.
definition category 1 1. Additionally.30–0.g. 1996.5 2 2. Sampling was interrupted for the duration of the recording in order not to overestimate locations in which animals had previously been captured. surface and depth. divided into seven categories corresponding to particular densities.5 m. for comparison of local with regional species pools).AMPHIBIAN MONITORING IN TROPICAL FORESTS turbance gradients (Jaeger 1994.35 m/s). In addition. Sampling methods and sampling effort Habitat characterization. For a discussion of the advantages of rectangular sample units over quadratic or circular ones see Krebs (1989) and McCune & Grace (2002). In order to quantify the availability of potential aquatic breeding sites.. Repeated controls of identical transects on consecutive days were avoided to ensure independence of samples. captured frogs were marked by TABLE 1. secondary forests.or age-specific differences in habitat use. A maximum of four transects was patrolled at night. We assumed that the number of plants with small dbh is greater in degraded. Pearman 1997). to assign them to particular guilds). In addition to species identity we recorded sex and snout-vent-length (SVL) of every individual. rectangular ones provide the possibility of easily up. thus providing data that are useful to test for sex. Usually four to six transects per day were sampled during daytime.g. These parameters included vegetation density in four strata (canopy: > 20 m.or downscaling data without neglecting local habitat diversity (e. Transects were intensively patrolled at a constant speed (0.5–1.. 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. Categories of habitat variables measured on transects. bush and shrub stratum: 0. Definitions of habitat variables are summarized in Table 1. 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. As far as possible all individuals were captured. and to investigate correlations of species assemblages with environmental variables. To avoid duplicate recordings.5 m). dbh).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 . whereas primary for- ests show increasing numbers of plants of larger dbh (Chatelain et al. Standardized visual transect sampling (SVTS). we characterized all habitats using several variables that were recorded at two defined points within each 25 m subunit (beginning and midpoint). 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. lower tree stratum: 3–10 m. thereby recording all amphibians within a distance of 100 cm from either side of the path. dbh = diameter of plants at breast height. substrate moisture was determined in four categories during every transect walk. understory: < 0. Edaphic parameters were registered as general substrate types according to Lieberoth (1982). To assess habitat preferences of particular species (e. Jaeger & Inger 1994). Specimens heard but not seen within this distance were not searched for and not included in the SVTS data sets (see below). Sampling was performed independent of prevailing weather conditions.5 3 3. Compared to linear transects. vegetation density was measured in four strata (see text). both in time and space.
habitat analyses can be based on data obtained from the general habitat characterization for single transects or segments. 1997). Anurans that are strong jumpers (e. Throughout transect walks a combination of visual and acoustic techniques was applied at all times. this technique allows the detecting of cryptic species that. The width of the acoustic transect depends on the ability to detect each species’ advertisement call. This device is useful to determine species richness of epigaeic organisms (Corn 1994). may be underestimated when exclusively using visual techniques. In the transect corners. This corresponds to 765 transect walks. In those cases. VES and AES are frequently used for rapid assessments and the evaluation of larger areas. Total VES. Furthermore. beginning with segment 1: segment 1. We used these methods continuously in all habitats of the study area including the transect areas. Calls from greater distances cannot be unambiguously identified and the chance of duplicate recordings in neighboring segments increases.. and to estimate relative abundances of species within an assemblage (Crump & Scott 1994). Visual (VES) and acoustic (AES) encounter surveys. thus creating 25 m x 25 m acoustic sampling plots. Likewise microhabitat description was not always possible.5 4 hours of transect sampling. despite their potential abundance. Drift fences consisted of durable green plastic gauze. Recaptures were excluded from the analyses. keeping the transect routine identical for both techniques. 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). VES can only provide information on the presence or absence of a species in an area but are inadequate for determining abundances. Individuals below nine mm SVL were not marked due to their small size. individual marking should be restricted to studies in which individual recognition is indispensable. during SATS it was often impossible to determine individual parameters. species composition. An area or habitat is searched systematically for individuals in a defined time period. and as the time-constrained searches of Corn & Bury (1990). 8.and AES-effort was kept comparable to the time spent on SVTS and SATS surveys throughout the entire study period. only calls coming from the right-hand side in the first segments were registered (i. In concordance with these authors the methods were only used as a qualitative or semi-quantitative tool within the scope of our studies. 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.e. as well as breeding habitat use and breeding phenology of species (Zimmerman 1994).. 0. 1994. Trapping with pitfall or funnel traps along drift fences. Bufo species). Dodd 1991). Capture success may vary greatly between species (Corn & Bury 1990. 12 and 20). Counts can be used to estimate relative abundance of calling males.g. 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. It can be used to determine the species richness of an area and the species composition of a local assemblage. Pitfall traps with drift fences were installed in two of the primary forest transects from March to April 1999. other than sex and species.RÖDEL & ERNST toe clipping (Donnelly et al. An array of fences and traps consisted of a central trap (buckets: .. This technique has been formalized as the time-constrained technique of Campbell & Christman (1982). Henle et al. The resulting data are expressed in numbers of individuals of a certain species found in an area per unit time.5 m to either side of the transect was defined. VES and AES thus could be used to evaluate if the randomly chosen transects represented the regional species pool. data can be expressed in numbers of individuals per time and surface units. For that reason a maximum recording distance of 12. Ptychadena species) are more difficult to trap than terrestrial species that lack these abilities (e.g. Coding schemes used for individual recognition were not applied. From February 1999 to December 2000 we compiled 382. For practical reasons “man-hours” can be used. As opposed to visual sampling. following the characterization routine used for general habitat description. which can be adjusted to the complexity of the habitats being sampled. and additionally in only one of these transects from January 2001 to September 2002. Audio strip transects represent a commonly used method for acoustic monitoring (Zimmerman 1994). Due to the simplicity of the method. Thus. For both SVTS and SATS. According to Corn & Bury (1990). At capturing sites a thorough description of the microhabitat was recorded.5 m high and stapled vertically onto wooden stakes. Standardized acoustic transect sampling (SATS).
Each segment was tightened around a plastic bucket with an opening angle of 45°. data). Pitfall trapping time summed to a total of approximately 4000 trap-days. 5 Ta ïF . The ends of each segment were flanked with additional plastic buckets. which are situated south and west of TNP respectively (Rödel & Branch 2002. unpubl. Additionally. and in a forest fragment (Paulé-Oula 2) outside TNP from 30 August to 12 September 2002 (A. SVTS = species recorded during visual transect walks. one on either side. fossorial and aquatic species independent of families. results were analyzed for leaf litter. TaïF = all species recorded with all methods in forest habitats around SRET station. SATS = species recorded during acoustic transect walks. Trap = pitfall and funnel traps. arboreal.. TC = species recorded during transect walks (SVTS and SATS combined). AES/VES = visual and acoustic encounter surveys (only species that have been recorded in forest habitats around SRET station). Duct tape was used to reduce the diameter of the buckets and construct funnel-like openings in order to impede escaping from traps. 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. we applied that method during a herpetological survey of the Haute Dodo and Cavally forests.AMPHIBIAN MONITORING IN TROPICAL FORESTS 275 mm deep. Numbers above bars represent total number of species recorded with the respective method. 285 mm top diameter. using a particular sampling method. Sampling success is referred to as number of species within families recorded. Traps were checked at least on a daily basis. Funnel traps summed to a total of 384 trap-days. Hillers et al. 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). 220 mm bottom diameter) and two triangular fence segments (total length 16 m). as sampling success and efficiency may vary with regard to the particular biology of a species. 1. Number of species per family recorded using different sampling methods. In addition. FIG. The data from pitfall and funnel traps can be expressed as number of individuals per trap-day. Branch & Rödel 2003). They were checked at least every morning.
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). Effectiveness in detecting species within given families differed significantly between methods (Friedman test. Hyperolius guttulatus). Rödel & Ernst 2000. VES/AES revealed 39 forest species (90. tested for TaïF. Taking all methods into account. bibroni. AES/VES. df = 5.g. 1.3 % of TNP species). p = 0. Three leaf litter species (Amnirana occidentalis. 1. 2).199. Appendix 1). SVTS.g. SATS. Phlyctimantis boulengeri ). H. fusciventris. Bufo superciliaris) or simply absence in the area under investigation (e.g. Seven species (Bufo maculatus. mascareniensis. TC. Four arboreal frogs were exclusively recorded with these methods (Hyperolius picturatus. B.0 % of TaïF) in eight families. 2. sp. N = 10.) were recorded exclusively outside forest habitats (e. thus making overall sampling success very high (Tab. Sampling success and data quality varied between different methods. and P. annulatus) were exclusively recorded during transect walks. RESULTS In total. Families containing only 6 a few species in the TNP area showed the highest recording rates. Ten species (23. Number of species per amphibian guild recorded using different sampling methods. clearing around SRET) and thus ignored in the analysis. rather than actual sampling method insufficiency.) or pitfalls only (Geotrypetes seraphini). regularis. and between different amphibian families or guilds (Figs. 2.RÖDEL & ERNST FIG. P. Rödel et al.7 %) of nine families. Phrynobatrachus fraterculus. compare Appendix 1). Failure to detect particular species in this region was most probably due to their general scarcity (e. During transect sampling (SVTS. Numbers above bars represent total number of species recorded with the respective method. χ2 = 19. Hoplobatrachus occipitalis..2 %) were recorded with pitfall and funnel trapping. 56 amphibian species are known to occur in TNP (Schiøtz 1967.002. Rödel 2000. Trap). nienokouensis. 2003. P.. Using all methods we recorded 50 species in the region of the SRET (89. Two fossorial species were exclusively detected either with VES and pitfall traps (Hemisus sp. Perret 1988. Ptychadena pumilio. 43 species were recorded in forest habitats within the study region (Fig. see Appendix 1). 1. H. SATS) we recorded a total of 15007 individual amphibians belonging to 37 species (86. P. . 2002. Phrynobatrachus calcaratus. For abbreviations see Fig..
VES and AES revealed highest species numbers.7 0.0 100.0 0. presence and abundance of purely aquatic species and tadpoles of all species were underestimated with all methods used here. 7 .0 100. A distance of 1 m (visual) and 12. asytlosternids) were best recorded with SATS. In TNP funnel traps were ineffective. For other abbreviations see text and Fig. As in TNP. 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). All other species were recorded with VES/AES (Rödel & Branch 2002.5 m (acoustic) proved to work equally well in all habitats investigated.0 100. unpubl.0 0.0 100. was difficult as detectability of species varied considerably with (e.0 0.4 0. tested for TaïF. SATS.2 66.03 specimens per trap-day in pitfall traps).0 SATS 0.0 100. p = 0.0 100. Branch & Rödel 2003).0 50. the fossorial Geotrypetes seraphini was recorded with pitfall traps only. in a primary forest transect with none in the forest fragment (Hillers et al.0 100. all were recorded using other methods as well. which is mute. N = 4. Reference is the number of species that were recorded using all methods in forested habitats around the SRET station (TaïF).0 0. neither between given family or guild.0 SVTS 0. Diurnal ground-dwelling frogs were best recorded quantitatively using SVTS. Family / guild Caecilidae Pipidae Bufonidae Hemisotidae Ranidae Petropedetidae Astylosternidae Arthroleptidae Hyperoliidae Rhacophoridae leaf litter arboreal fossorial aquatic TaïF 1 1 2 1 5 11 1 3 17 1 22 18 2 3 TC 0.0 36.0 31..0 0. In contrast.0 100. transect walks made it possible to search for amphibians in a very comparative way. However. Most arboreal frogs could be recorded with SVTS. We only captured a few leaf litter frogs. With the exception of Acanthixalus sonjae.0 100.0 100.0 66. 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. but very low in arboreal species (6. data).0 Trap 100.5 100.294.5 % of recorded leaf litter frogs). quantitative data for all arboreal species were best assembled with SATS.8 100.0 100.0 40.0 76. df = 4. 1990). other than based on time units.7 100.47 specimens per trap-day in contrast to 0.0 81.5 100.0 100. nocturnal leaf litter frogs (arthloeptids.0 64.g.) vegetation density.0 77. SVTS.0 100. Schaich & Hamerle post hoc multiple comparisons revealed no significant differences between methods at α = 0.0 72.0 80. No additional species were detected with funnel traps. 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 50.0 100.AMPHIBIAN MONITORING IN TROPICAL FORESTS TABLE 2.0 76. Trapping success was highest in leaf litter frogs (56.05.0 100.0 0.067.0 100.0 AES/VES 0. Percentage of species within an amphibian family or guild that were recorded using a particular method.0 77. VES/AES.0 100.0 100.0 100.8 0.0 100.0 0.0 0.0 100.0 100. but that concerned mostly single specimens.8 0. Quantification of these data.3 We could not detect a statistical difference in sampling efficiency between different amphibian guilds using the methods employed (Friedman test. 1.7 % of recorded arboreal species).7 0.8 0.0 100. Most specimens were captured with funnel traps (0.0 86.4 100.7 100. Trap).0 100. χ2 = 10. TC. when being a part of the transect line itself.0 68. Likewise.0 100. As aquatic sites were only taken into account.0 33.0 100.0 100.
but appear not to be appropriate when standardized quantitative sampling is required. The advantage of transect sampling was especially obvious when focusing on leaf litter species. such as species groups. Migrating pipid frogs can also be easily recorded with pitfall and funnel traps. especially arboreal and fossorial. such as the Hylidae. iii: efficiency. can then be compared to the real conditions in an area or habitat. Although widely used and recommended. We found it impossible to quantify VES and AES other than in relation to time. The combined transect sampling methods provided a close approximation to the real presence and abundance of species. e. whereas visual sampling appeared to be superior to acoustic sampling. which are mute (Drewes 1984. Methods of collecting quantitative data for aquatic amphibians and for tadpole assemblages have been described by Shaffer et al. When including relative abundance measurements these methods were insufficient. Olson et al. such as the very abundant Arthroleptis sp. 1996).1. these methods do not seem to sample a given fauna adequately well (Allmon 1991. Bury & Corn 1987. In this study this has been done by determining relative species richness with reference to families or guilds. a combination of SVTS and SATS. Therefore transect sampling can be considered to be the method of choice when sampling leaf litter anurans.2. and iv: environmental impact. Simple presence/absence data of aquatic species and stages can best be gathered by direct observation (surfacing individuals) and with dip-netting. (1994). Rödel & Ernst 2003). In addition. this only holds true if species numbers alone are considered. as an adequate measure. acoustic sampling appeared to be indispensable since these species were more readily detected by their calls than 8 by sight. during rapid assessment surveys (for recent West African examples see Rödel & Branch 2002. 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.RÖDEL & ERNST DISCUSSION Choice of methods should reflect the best compromise between i: practicability. Cardioglossa leucomystax. Whereas transect walks provide the possibility of monitoring amphibians in a very comparative way. Rödel 2003. They suggest using relative species richness within taxonomic entities. We found no additional species and only very few specimens by funnel trapping. Although the combination of AES and VES provided an adequate approximation to real conditions (see also Doan 2003).g. were nearly as successful in detecting species. This was also true for most of the arboreal species belonging to the families Hyperoliidae and Rhacophoridae. VES and AES yielded highest species numbers. Arthroleptis sp. especially in long-term field studies within temperate regions (e. as they impose a severe disturbance on the system under investigation and thus were not suitable to our goals. Pitfall trapping was only useful in detecting fossorial species that were not encountered during transect walks. inaccessible areas than in areas that are easy to monitor. or Phrynobatrachus alticola. According to Pearman et al. in order to sample particularly secretive leaf litter species. We found it more realistic to apply a (clearly subjective) measure for habitat complexity on which to base collection effort during VES/AES. SVTS was generally the best method for detecting diurnal leaf litter frogs. 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. Rödel et al. as well as SVTS alone. Doan 2003). VES and AES are useful tools for the compilation of species inventory lists.. 2003). and failed to detect aquatic and fossorial species. determined by using a particular sampling method. The differences in efficiency of visual versus acoustic sampling varied between taxa.g. Semlitsch et al. However. In general SATS was very efficient in detecting nocturnal leaf litter frogs and treefrogs but less successful in detecting diurnal leaf litter frogs. Vonesh 2001. pitfall traps and drift fences . Some taxa. could be detected exclusively using visual sampling methods. The methods VES and AES in combination with pitfall traps with drift fences were useful in sampling additional.. but is certainly just as well suited for the investigation of tropical anuran assemblages in general. genera or families. The percentage of species within supraspecific taxonomic units. such as species of the genus Acanthixalus. Standard plot sampling and total removal plots were not tested. ii: specific aims. Schiøtz 1999. However. species not encountered during transect sampling. (1997) and Rödel (1998a). (1995) a simple comparison of species numbers may not be sufficient to evaluate the efficiency of different sampling methods. Thus even time-based quantification of these data is questionable. spending more time in complex. The methods tested within this study proved to perform with varying degrees of efficiency. Nonetheless.
In our experience. (2004) had high trapping success with pitfalls in Gabon when considering numbers of individuals. Thus data should be gathered throughout a whole season to obtain a thorough knowledge of the local fauna. Burger et al.6 % of all individuals captured with this method. amphibian guild and environmental impact. With the exceptions of aquatic and fossorial species they pro- vided data that cover the whole community very well. traps have to be uninstalled or closed and reopened when starting again. this method cannot be recommended for standardized sampling of tropical anurans.g. be quantified in relation to time. in theory.g. If the investigation is interrupted for some time. QS/TR = quadrat sampling/total removal sampling (data from Allmon 1991. as well as environmental changes. A comparison of trapping data with data other than those of the observer in question seems to be difficult. SVTS and SATS provided data on species and habitats und thus may reveal changes in composition and species abundance. VES and AES are less time consuming. Theoretically it might well be that trapping will result in dislocating at least some species from the area where traps have been established. Rodda et al. experience of investigator and prevailing weather conditions (see Branch & Rödel 2003).AMPHIBIAN MONITORING IN TROPICAL FORESTS proved to be the least effective method. Transect walks are time intensive. Costs with regard to time spent for data acquisition was highest in trapping and in transect walks. in consecutive field seasons. Effectiveness of methodology in regard to type of data required. (2001) report on similar experiences using this method in a herpetological survey in Guyana. the fossorial Hemisus perreti and the aquatic Silurana epitropicalis. 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 . Donnelly et al. They collected only six species (out of a total of 132 species of amphibians and reptiles being recorded). e. however. accounted for 72. if not on amphibians then on other organisms. However. Doan 2003). Data can be used for comparisons between habitats. one should test the efficiency at a particular site in advance before installing these traps on a broad scale. the frequency of sampling can be reduced when general phenological traits of the species recorded have been clarified. seasons and years.. Vonesh 2001. for other abbreviations see text and Fig. as trapping success per trap per time unit. However trapping success depends very much on choice of site. SVTS and SATS have been proven to be the only methods that provided quantitative data on forest amphibians with regard to space and time. Parris 1999. generalization of data with regard to phenology or abundances is less possible. Data from both trapping methods can. It is advisable to perform them randomly independent of prevailing weather conditions.) that leopard (Panthera pardus) tracks TABLE 3. When considering the relatively high costs and time investment due to the high level of maintenance required (Parris 1999). We observed (e. Transects will have an environmental impact. 2001. The impact of traps and drift fences was difficult to judge. However only two species. 1. VES and AES had probably the lowest environmental impact. Traps have to be checked at least daily. none of them uniquely found by this method.
Branch.. Glos.. 10 . Pp. Memoir 28: 145–186. 5: 37–53. 2004. Observations sur la reproduction et la dynamique des populations de quelques anoures tropicaux I. and C. L.-A.P. Standard methods for amphibians. in the worst case.. M. Christman. Donnelly. Amphibians and reptiles of Monts Doudou. 1998. & M.o. A. Bull. G. noire. Wildlife Research Report 13. 2 ed. 1997. P. possibly accompanied by opportunistic trapping. 30: 133–165. Ecol. Phrynobatrachus accraensis. Trefaut Rodrigues.R. Channing... Sci. C. 1979b.. consequent long-term effects on forest structure. However. fond. R. 1994. Trefaut Rodrigues. & M. 193–200 in Scott. 2003) transects should be supplemented by specifically investigating aquatic breeding sites with the appropriate set of standardized techniques (Heyer et al.S.R. 1964. Gbamlin.S.E. 1996...M. W. Bortz. Department of the Interior. 2003. Campbell. Corn.. Rödel. Conserv. Biodiv. not only on their breeding sites but throughout the whole range of habitats used by them. Sér..S.. The “Projet Autonome pour la Conservation du Parc National de Taï”. Richards. 01 LC0017). Transect cutting and the use of transects should therefore be done with the necessary precautions. Basset. Barbault. Hammond.. Pp. Ptychadena maccarthyensis et Ptychadena oxyrhynchus. J. 1991.S. R. J. Research permission was given by the “Ministère de l’Enseignement Supérieur et de la Recherche Scientifique..J. L. Field techniques for herpetofaunal community analysis. Lienert. Barbault. Wildlife Management 51: 112–119. 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. Observations sur la reproduction et la dynamique des populations de quelques anoures tropicaux III. N’Dri. Bury.. & S. W. N. Central Amazon. Spichiger. Inst. Project W08 BIOTA-West.1964. Braun-Blanquet. M. Straight-line drift fences and pitfall traps. TROPENBOS . ACKNOWLEDGMENTS This study is part of the BIOLOG-program of the German Ministry of Education and Science (BMBF. Alford. for specific questions (cf. Hayek.. Gautier. Terre Vie 32: 441–452. A recent history of forest fragmentation in southwestern Ivory Coast. Herpetological survey of the Haute Dodo and Cavally forests. Hofer.A. Calif. 1978. Y. Pflanzensoziologie. & P. (ed.. Washington & London. 7: 503–522. & R. Canopy fogging of an overstory tree – recommendations for standardization. R. 1987. J.W. Afr.-O.Côte d’Ivoire helped with transportation. Berlin a.W.” G... U.Y.G. (eds. Boehnke.RÖDEL & ERNST were more often seen on the transect path than in undisturbed forest. Lips et al. Olson et al. A plot study of forest floor litter frogs.. Furthermore transects provide an effective method of investigating leaf litter frogs in particular. Observations sur la reproduction et la dynamique des populations de quelques anoures tropicaux IV. western Ivory Coast. & K.. A 41: 417–428. W. 865 pp. 1999. Wien. W. Corn. & S.J. Verteilungsfreie Methoden der Biostatistik.A. H. a well visible path becomes established that at least in primary forests remains so for years. 1990. Syst. R. Trefaut Rodrigues. 1979a. Ann. Acad. 109–117 in Heyer. The presented transect design proved to be adequate for representative sampling and appeared to be appropriate for most studies involving multivariate structure data. REFERENCES Adis.. Springer Verlag.. J. Jr. After using transects intensively for some time. Chatelain.): Herpetological communities. Part II: Trapping results and reptiles. J. Linsenmair. 3. Hillcoat provided valuable comments on the manuscript. Brazil. Foster. Ecol. & K. For simple short-term surveys we recommend VES and AES. 20: 64–77.. SVTS and SATS are especially useful for long-term studies. R. McDiarmid. Arthroleptis poecilonotus. 1982. Rev. This might affect densities of other mammals including possible seed dispersers with. Barbault. and the “Taï Monkey Project” provided logistic support. Floren.” of the Republic of Ivory Coast. Allmon. Evaluation of pitfall trapping in northwestern forests: trap arrays with drift fences.” Working in TNP was kindly permitted by the Commandant K. Gabon: Species turnover along an elevational gradient. thus generating a much more complete picture of an amphibian community than is possible with other methods. Halliday. Ecol. Springer Verlag. Trop. Fish and Wildlife Service 13. & M. and B.): Measuring and monitoring biological diversity. & M. 364 pp. & A. Trop. Salamandra 39: 91–110. T. The access permit to TNP was issued by the “Ministère de la Construction et de l’Environnement.D. 1994. P. Global amphibian declines: a problem in applied ecology. Ecotropica 4: 93–97. J. Grundzüge der Vegetationskunde. 939 pp. Rödel 1998a). & M. R.A.C. Smithsonian Institution Press. U. Branch.R. All this support is gratefully acknowledged. Advantages and disadvantages of different methods are summarized in Tab.B. Ouoro provided invaluable help during fieldwork. Burger.
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3 = Phrynobatrachus guineensis is a leaf litter frog but reproduces in small water-filled tree holes (Rödel 1998b). TaïF = species recorded in forest habitats around the SRET station. Perret 1988) but not recorded by us. TC = all species recorded on transects. 6 = recorded north of area under investigation.AMPHIBIAN MONITORING IN TROPICAL FORESTS APPENDIX. 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. 5°25’ N. other than transects. TS = all species recorded on transects in secondary forest. 5 = treated as full species and not as subspecies of Hyperolius fusciventris due to sympatric occurrence in TNP (see Rödel & Branch 2002). 2 Ranidae Hoplobatrachus occipitalis Amnirana albolabris Amnirana occidentalis Aubria occidentalis Ptychadena pumilio Ptychadena bibroni Ptychadena mascareniensis Ptychadena superciliaris Ptychadena aequiplicata Ptychadena longirostris Ptychadena sp. a l l l l l 13 . 2 = determination not possible at present time. species Gymnophiona Caecilidae Geotrypetes seraphini Anura Pipidae Silurana tropicalis Bufonidae Bufo regularis Bufo maculatus Bufo togoensis Bufo taiensis Bufo superciliaris Hemisotidae Hemisus sp. 1 = known from TNP (Schiøtz 1967. List of amphibian species recorded in Taï National Park (TNP). f = fossorial species. 2003). a = arboreal species. 4 = Acanthixalus sonjae lives aquatically in large water filled tree holes but forages in surrounding vegetation (Rödel et al. SRET = species recorded with VES/AES on the clearing around the SRET station. 7°10’ W). Trap = species recorded with pitfall and funnel traps. G = species recorded in southern TNP (Guiroutou. VES/AES = species recorded via visual and acoustic encounter surveys in forest habitats near SRET. aq = aquatic species. TP = all species recorded on transects in primary forest. l = leaf litter species.
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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