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339905541-Oa unsuitable for oviposition by S. albicans. However, no significant Herpetological Review, 2006,37(2), 161-165. negative relationship was recorded. (0 2006 by Society for the Study of Amphibians and Reptiles The patterns found show aggregation size to be remarkably con- stant by the standards of any vertebrate populations. The perma- Abundance and Biomass of Twelve Species of nence of males in aggregation was high, and the little variation in Snakes Native to Northeastern Kansas aggregation size was explained by variations in the numbers of males entering the aggregation each month. The possibilility that HENRY S. FITCH aggregation size could be used as in index to population size is University of Kansas Fitch Natural History Reservation jeopardized by its high constancy and lack of relationship with 2060 East 1600 Road, Lawrence, Kansas 66044, USA rainfall patterns. Possibly aggregation size is rather controlled by social factors regulating the number of males calling at a time in and ALICE F. ECHELLE any given place. Department of Zoology, Oklahoma State University Stillwater, Oklahoma 74078, USA Acknowledgments. —We thank IBAMA (Institute Brasileiro de Meio e-mail: [email protected] Ambiente e Recursos Naturais Renovaveis) for allowing the work at Parque Nacional da Serra dos Orgaos; Institute Nacional de Meteorologia The purposes of this paper are to examine abundance and biom- (INEMET) for climatic data; Sergio Potsch de Carvalho e Silva for teach- ass estimates of the dozen most abundant snake species in our ing on amphibians, for logistic support and co-supervising DN's MSc area of northeastern Kansas, and to relate these numbers to infor- dissertation; the several colleagues, especially Fabio Nascimento, who mation reported in the literature. Hirth and King (1968) stated, "In helped in field work; Romulo Barroso for company in all field work and spite of all the current interest in ecosystem ecology there is still a discussions throughout the study; Jose P. Pombal Jr., William Magnusson, dearth of information concerning biomass densities of snakes in Jon Loman, Monique Van Sluys, Ronaldo Fernandes, Gunther Koehler and an anonymous referee for very useful comments on the manuscript; various habitats." Except for a few studies (e.g., Bonnet et al. 2002; Jorge Luiz do Nascimento for ideas in this study and for financial sup- Godley 1980; Hirth and King 1968; Reichenbach and Dalrymple port. 1986; Winne et al. 2005), the situation does not seem to have changed appreciably in the last 37 years. LITERATURE CITED METHODS AND STUDY AREA BOKERMANN, W. C. A. 1967. Dos nuevas especies de Hyla del grupo catharinae (Amphibia, Hylidae). Neotropica 13:62-66. Snakes have been studied for the past 56 years on Kansas DEAN, W. 1996. A Ferro e Fogo – A Historia e a Devastacao da Mata Atlantica Brasileira. Companhia das Letras, Sao Paulo. 484 pp. University's Natural History Reservation (FNHR) following es- FERNANDEZ, F. A. S. 1995. Metodos para estimativa de parametros tablishment of the area as a reserve. Although protected from an- populacionais por captura-marcacao-recaptura. Oecologia Brasiliensis thropogenic alterations, the area has undergone continual and pro- 2:1-26. gressive change due to natural ecological succession (Fitch 1999; HEYER, W. R., A. S. RAND, C. A. G. CRUZ, AND 0. L. PEIXOTO. 1988. Deci- Fitch et al. 1984, 2001, 2003b). The closing in of forest and elimi- mations, extinctions, and colonizations of frog populations in South- nation of open places associated with agriculture and grazing has east Brazil and their evolutionary implications. Biotropica 20:230-235. been unfavorable for snakes. Several of the species have disap- HOULAHAN, J. E., C. S. FINDLAY, B. R. SCHMIDT, A. H. MEYER, AND S. L. peared from this 239-hectare tract, and almost all species have Kuzmusr. 2000. Quantitative evidence for global amphibian population declines. Nature 404:752-755. been drastically reduced (Fitch 1999). The adjoining Nelson En- MARTOF, B. S. 1953. Territoriality in the green frog, Rana clamitans. Ecol- vironmental Study Area (NESA), acquired by the University in ogy 34:165-174. 1970, has provided a contrast to FNHR in terms of habitat stabil- NASCIMENTO, D. 2003. Comportamento reprodutivo de Scinax albicans ity. Invasion of woody vegetation is prevented on NESA by the (Bokermann, 1967) (Amphibia, Anura, Hylidae), na floresta pluvial regular mowing of blocks of former pasture and cultivated land, montana no sudeste do Brasil. Unpubl. M.Sc. Dissertation, Museu and as a result, existing snake populations resemble those found Nacional, Rio de Janeiro. on FNHR in earlier stages of succession (Fitch 1999). PECHMANN, J. H. K., D. E. SCOTT, R. D. SEMLITSCH, J. P. CALDWELL, L. J. Sampling areas (House Field, Quarry Field, etc.) are described Vrrr, AND J. W. GIBBONS. 1991. Declining amphibian populations: the by Fitch (1999). Of the 12 censuses reported here, eight were from problem of separating human impacts from natural fluctuations. Science 253:892-895. FNHR and were done in the earlier years of study (before 1979) SEBER, G. A. F. 1982. The Estimation of Animal Abundance and Related except that for Thamnophis sirtalis. Of the remaining four spe- Parameters. 2nd ed. Charles Griffin, London. 654 pp. cies, Carphophis vermis was censused by Clark (1970) on private WEYGOLDT, P. 1989. Changes in the composition of mountain stream frog property adjoining FNHR and three, Lampropeltis triangulum, L. communities in the Atlantic Mountains of Brazil: frogs as indicators of calligaster, and Coluber constrictor, were censused post-1990 on environmental deteriorations? Stud. Neotrop. Fauna Environ. 243:249- NESA. 255. Unless otherwise noted, density estimates are based on field YOUNG, B. E., K. R. LIPS, J. K. REASER, R. IBAREZ, A. W. SALAS, J. R. records and censuses originally published in Fitch (1999) except CEDERO, L. A. COLOMA, S. RON, E. LA MARCA, J. R. MEYER, A. MuSroz, for Clark's (1970) Carphophis figures. Mark-recapture records and F. BOLAS1OS, G. CHAVES, AND D. Rollo. 2001. Population declines and priorities for amphibian conservation in Latin America. Cons. Biol. the Petersen Index were the bases for the abundance estimates for 15:1213-1223. most species, including Agkistrodon contortrix (Copperhead), ZAR, J. H. 1999. Biostatistical Analysis. 4th ed. Prentice Hall, Englewood Carphophis vermis (Western Wormsnake), Coluber constrictor Cliffs, New Jersey. 663 pp. (Racer), Crotalus horridus (Timber Rattlesnake), Diadophis Herpetological Review 37(2), 2006 161 punctatus (Ring-necked Snake), Nerodia sipedon (Northern of C. horridus from FNHR was in 1964. By contrast, 12 were Watersnake), Pantherophis obsoletus (Black Ratsnake), Pituophis captured from a 10-ha area of NESA (Biotic Succession Area) in catenifer (Bullsnake), and Thamnophis sirtalis (Common the years 1990 through 2002 (Fitch et al. 2003). Additionally, in Gartersnake). For the remaining three species, Lampropeltis the spring of 2003, 26 were captured along a rock outcrop about calligaster (Prairie Kingsnake), Lampropeltis triangulum 100 m long near Frank B. Cross Reservoir, in an area adjoining (Milksnake), and Storeria dekayi (DeKay's Brownsnake), densi- NESA on the east (Fitch et al. 2004). ties were estimated from their respective numbers relative to one Diadophis punctatus-Many different censuses are available or more species with larger mark-recapture samples. for the once-abundant Ring-necked Snake (Fitch 1999). Seven were Weights are based on figures from Fitch (1999, Table 98). Num- done in Quarry Field in 1966-67 and 1969-70, with sampling pe- ber of snakes, weight, and biomass per hectare were log trans- riods ranging from 0.5 to 2 months Eliminating the extreme low- formed for the regressions. Representative weight for each spe- est and highest estimates (597 and 4000/ha) gave a range of 791- cies is the midpoint between the means for the adults of both sexes 2039/ha and a mean of 1325/ha. An additional eight censuses were (excluding gravid and recently parturient females). For the sake from House Field with sampling periods ranging from 0.5 to 3 of consistency, only adult mass was used to calculate biomass from months (1965-1967); excluding the lowest and highest estimates density, although immatures were present for every species (262 and 1792) gave a mean of 1224/ha (range = 773-1761). Av- censused (see Fitch 2000, for age pyramids for 10 of the species). eraging the means for these two areas, both on FNHR, gives an Attempting to factor in mean mass across all sizes had the poten- overall mean of 1275/ha. tial of introducing more bias than was inherent in excluding these Lampropeltis calligaster-2.5/ha, 1990-1997, on the northwest- other size classes. Relative abundance of life history stages varied ern pens area of NESA. This estimate is the average of 1.6/ha markedly over each trapping season, but these seasonal fluctua- (based on ratio of capture records compared to Thamnophis) and tions were not synchronized among species. Also, age composi- 3.3/ha (compared to Coluber). Since 2001, 14 L. calligaster have tion of each species censused was partly a function of trapping been captured on NESA, and only two individuals (both in 2002) method. For example, entrance holes of the wire funnel traps were have been found on FNHR. mostly too small to allow entry of large adult Timber Rattlesnakes. Lampropeltis triangulum-0.6/ha, 1990-1997, on the north- In contrast, the traps were well suited to catch larger young and western pens area of NESA. This estimate is the average of 0.52/ adults of the Common Gartersnake, while tiny neonates of this ha (compared to Thamnophis) and 0.73/ha (compared to Coluber). species easily passed through the quarter-inch mesh of the traps, The last capture of L. triangulum on FNHR was in 1999. Since and hence are missing from the trapping records. More informa- 2001, 17 have been captured on NESA.
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