––––––, R. MATHEWS, AND R. KINGSINGER, JR. 1964. The mouth parts oftadpoles of Hurter’s spadefoot. Herpetologica 19:284–285.BROWN, H. A. 1989. Tadpole development and growth of the Great Basinspadefoot toad, Scaphiopus intermontanus, from central Washington.Can. Field Nat. 103:531–534.CRUMP, M. L. 1986. Cannibalism by younger tadpoles: another hazard ofmetamorphosis. Copeia 1986:1007–1009.DURHAM, F. E. 1956. Amphibians and reptiles of the North Rim, GrandCanyon, Arizona. Herpetologica 12:220–224.GOSNER, K. L. 1960. A simplified table for staging anuran embryos andlarvae with notes on identification. Herpetologica 16:183–190.HALL, J. A. 1993. Post-embryonic Ontogeny and Larval Beahvior of theSpadefoot toad, Scaphiopus intermontanus (Anura: Pelobatidae). Ph.D.thesis, Washington State Univ., Pullman.––––––. 1998. Scaphiopus intermontanus Cope Great Basin spadefoot.Cat. Amer. Amphib. Rept. 650.1–17.––––––, J.H. LARSEN JR, AND R.E. FRITZNER. 1997. Postembryonic ontogenyof the spadefoot toad, Scaphiopus intermontanus (Anura:Pelobatidae): external morphology. Herpetol. Monogr. 11:124–178.––––––, ––––––, AND ––––––. 2002. Morphology of the prometamorphiclarva of the spadefoot toad, Scaphiopus intermontanus (Anura:Pelobatidae), with an emphasis on the lateral line system and mouthparts.J. Morphol. 252:114–130.HAMPTON, S. H., AND E. P. VOLPE. 1963. Development and interpopulationvariability of the mouthparts of Scaphiopus holbrooki. Amer. Midl.Nat. 70:319–328.MAYHEW, W. W. 1965. Adaptations of the amphibian, Scaphiopus couchi,to desert conditions. Amer. Midl. Nat. 74:95–109.MOREY, S., AND D. REZNICK. 2004. The relationship between habitat permanenceand larval development in California spadefoot toads: fieldand laboratory comparisons of developmental plasticity. Oikos104:1736–1749.NUSSBAUM, R. A., E. D. BRODIE, JR., AND R. M. STORM. 1983. Amphibiansand Reptiles of the Pacific Northwest. University Press of Idaho, Moscow,Idaho.ORTON, G. L. 1954. Dimorphism in larval mouthparts in spadefoot toadsof the Scaphiopus hammondii group. Copeia 1954:97–100.PFENNIG, D. W. 1990. The adaptive significance of an environmentallycueddevelopmental switch in an anuran tadpole. Oecologia 85:101–107.––––––. 1992. Proximate and functional causes of polyphenism in ananuran tadpole. Functional Ecology 6:167–174.POMEROY, L. V. 1981. Developmental polymorphism in the tadpoles ofthe spadefoot toad Scaphiopus multiplicata. Ph.D. thesis, Univ. of California,Riverside.POTTHOFF, T. L., AND J. D. LYNCH. 1986. Interpopulation variability inmouthparts of Scaphiopus bombifrons in Nebraska (Amphibia:Pelobatidae). Prairie Nat. 18:15.STORZ, B. L. 2004. Reassessment of the environmental mechanisms controllingdevelopmental polyphenism in spadefoot toad tadpoles.Oecologia 141:402–410.TANNER, V. M. 1939. A study of the genus Scaphiopus: the spadefoot toads.Great Basin Nat. 1:3–19.TURNER, F. B. 1952. The mouth parts of tadpoles of the spadefoot toad,Scaphiopus hammondii. Copeia 1952:172–175.<strong>Herpetological</strong> <strong>Review</strong>, 2008, 39(2), 154–155© 2008 by Society for the Study of Amphibians and ReptilesSpring Peepers and Pitcher Plants: A Case ofCommensalism?RONALD W. RUSSELLDepartment of Biology, Saint Mary’s University923 Robie Street, Halifax, NS, Canada, B3H 3C3e-mail: ron.russell@smu.caSarracenia purpurea (Northern Pitcher Plant) is a carnivorousplant found throughout northeastern North America (Schnell 2002).This plant is frequently encountered in nutrient-poor bogs, oftenassociated with Sphagnum spp. The water-filled, pitcher-shapedleaves of S. purpurea serve as a trap for small invertebrates attractedby extra-floral nectaries near the entrance to the pitcher.Once entry to the pitcher is accomplished, escape is difficult dueto downward oriented hairs on the inner surface of the leaf, andcaptured organisms drown in accumulated rainwater. Nutrientsfrom decomposing invertebrates are absorbed by the plant (Ellisonand Gotelli 2001). Capture efficiency of insect prey in NorthernPitcher Plants is low (0.83–0.93%) (Newell and Nastase 1998). Amutualistic relationship is hypothesized to exist between pitcherplants and the inquiline community contained within the pitchers(Bradshaw and Creelman 1984; Ellison and Gotelli 2001). Tinyvertebrates are also known to become entrapped in Sarraceniapitchers (Schnell 2002).There are a number of anecdotal references to amphibian consumptionby Sarracenia pitchers as well as pitcher use by amphibiansin the popular press (The Sentinel 2007). One of the earliestreferences to Sarracenia describes pitchers as insect refugiafrom amphibian predation (Catesby 1743). Amphibians are knownto become entrapped and digested in pitchers (Butler et al. 2006;Schnell 2002), forage for insect prey on pitchers (Jones 1935),and inhabit pitchers (Lim and Ng 1991). In this study, I quantifypitcher use by Pseudacris crucifer (Northern Spring Peeper) andelucidate the nature of the frog-pitcher plant interaction.Twelve adult Northern Spring Peepers (8 females, 4 males) werecollected from the field in early May 2004 and placed in a 90-literglass terrarium extensively planted with Sphagnum sp. and fourNorthern Pitcher Plants with 5–11 pitchers per plant. Pitcher plantdensity in the laboratory was similar to plant densities observed inthe field and peepers had access to non-dessicating roosting siteswithin the Sphagnum mat. All spring peepers were reproductiveand ranged in SVL from 18–26 mm. Spring peeper density in thelaboratory was much greater than observed in the field. Amphibianswere fed wingless fruit flies and juvenile crickets. The artificialhabitat was observed at least 3 days per week from May–September for 15 minutes per day. Fruit flies were attracted toextra-floral nectaries on pitchers and spring peepers were frequentlyobserved (at least once per observation period) climbing pitchersto consume these insects. Peepers were routinely observed insideS. purpurea pitchers during the day, but were never observed feedingwhile inside pitchers. Suitably sized pitchers of all plants wereoccupied and no territorial behavior was observed. Occupancy rateswere typically less than 5% (0, 1, or 2 peepers observed in pitchers).Only pitchers large enough to admit peepers were used to154 <strong>Herpetological</strong> <strong>Review</strong> 39(2), 2008
estimate occupancy. The entrance to the smallest pitchers was toosmall to admit adult frogs, but could potentially accommodate recentlymetamorphosed spring peepers. This microcosm was maintainedfor six months, with no amphibian losses resulting fromentrapment in pitchers.Sarracenia purpurea plants located on a coastal barren nearHalifax, Nova Scotia, Canada (44°33.246'°N, 63°31.396'W) weresurveyed for the presence of spring peepers. In 2004, 163 pitcherswere observed and 306 in 2005. Density of Northern Pitcher Plantsbased on ten random 1 m quadrats was 8.5 ± 4.4 plants/m 2 . Quadratswere selected by overlaying a sketch of the study area usingapproximate distances with a grid, randomly numbering the squaresof this grid, then selecting ten squares by a random number generatorfor density estimates. Spring peeper surveys were conductedover three consecutive days during late May in both years in alarge bog dominated by Sphagnum sp., Eriophorum sp., and Sarraceniapurpurea where plants grew on a nearly continuous matof Sphagnum. Observations were limited to a relatively short periodat the peak of the spring peeper breeding season since theseamphibians disperse back into the forest following reproduction.Additionally, in late May 2005, 50 pitchers were randomly collectedfrom this site, dissected by longitudinal incision with a scalpelin the laboratory, and examined for the presence of amphibianremains. Five pitchers of sufficient size were collected from tenquadrats, randomly selected as described above.There were no peepers observed in Sarracenia pitchers in 2004,however in 2005, four spring peepers were observed in pitchersduring daylight. This corresponds to an incidence of 1.3% of pitchersoccupied. When a pitcher was determined to be occupied by aspring peeper, it was marked with a ring of jute twine. Markedpitchers were inspected the following day and it was observedthat none was occupied on consecutive days, indicating that amphibianshad moved prior to the second survey. Frogs were nottrapped within the previously occupied pitchers. Night surveys of12 wetlands on this same coastal barrens from 2002 to 2006 (atleast 8 surveys per year) revealed a large and active spring peeperpopulation based on call surveys. Peepers were occasionally observedcalling from pitchers at night and none of the 50 dissectedpitchers contained identifiable amphibian remains. Complete digestionfor trapped amphibians in S. purpurea pitchers requires atleast 10 days (Butler et al. 2006). The sample of dissected pitcherswas probably too small to detect amphibian remains with 1% orless.The combination of laboratory behavior and field observationsindicate that spring peepers occasionally use Sarracenia pitchers.Peepers forage for small insects attracted by extra-floral nectarieson the pitcher, intercepting invertebrates that might otherwise becomeplant food. There is no obvious advantage to the plant fromthis interaction, thus no mutualistic association, as observed withthe pitcher-inquiline community, which assists in decompositionand release of nutrients to the plant. By intercepting nutrition thatwould normally be routed to the plant and inquiline community,spring peepers may function as parasites; however, only a smallfraction of insects attracted to the extra-floral nectaries becomeentrapped in the pitcher (Newell and Nastase 1998). Parasitism isa minor interaction because of the low incidence of peepers inpitchers. Peepers may function as commensals by harvesting insectsattracted to the pitcher.Spring peepers forage for insect prey mostly during the day(Oplinger 1967), thus exposing these amphibians to potentiallydesiccating conditions. Peepers avoid desiccation during dry conditionsby moving under debris (Wright and Wright 1949). Themoist environment of the Sarracenia pitcher provides an ideal refugefrom desiccation for amphibians. While refuge in the pitchermay appear to be neutral to the plant, the amphibian partially occludesthe entrance to the digestion chamber by taking residencein the pitcher, which may affect pitcher trapping ability. Additionally,the frog is in an ideal position to consume trapped insects.This interaction clearly does not benefit the plant. Use of pitchersas a refuge was a rare event (1.3%) at the study site therefore thiswas not an important intertaxa interaction at this particular location.The importance of this interaction may increase where pitcherplants are less abundant. While use of Northern Pitcher Plants byspring peepers is not beneficial to the plant, it is a rare event. Consumptionof amphibians by pitcher plants is an equally rare event(Butler et al. 2006) which could compensate plants for nutritionusurped by spring peepers.Acknowledgments.—Funding was provided by a Discovery Grant fromthe Natural Sciences and Engineering Research Council of Canada. Permitswere obtained from the Nova Scotia Department of Natural Resourcesand the local Animal Care Committee. I thank A. M. Ellison and an anonymousreviewer for constructive comments on this manuscript.LITERATURE CITEDBRADSHAW, W. E., AND R. A. CREELMAN. 1984. Mutualism between thecarnivorous purple pitcher plant and its inhabitants. Am. Midl. Nat.112:294–304.BUTLER, J. L., D. Z. ATWATER, AND A. M. ELLISON. 2006. Red-spotted newts:an unusual nutrient source for northern pitcher plants. Northeast. Nat.12:1–10.CATESBY, M. 1743. The Natural History of Carolina, Florida, and theBahama Islands, Vol. II. Benjamin White, London, England.ELLISON, A. M., AND N. J. GOTELLI. 2001. Evolutionary ecology of carnivorousplants. Trends Ecol. Evol. 16:623–629.JONES, F. N. 1935. Pitcher plants and their insect associates. In M. V.Walcott (ed.), Illustrations of North American Pitcher Plants, pp 25–34. Smithsonian Institution Press, Washington, DC.LIM, K. K. P., AND P. K. L. NG, 1991. Nepenthiphilous larvae and breedinghabits of the sticky frog, Kalophrynus pleurostigma Tschudi (Amphibia:Microhylidae). Raffles Bull. Zool. 39:209–214.NEWELL, S. J., AND A. J. NASTASE. 1998. Efficiency of insect capture bySarracenia purpurea (Sarraceniaceae), the northern pitcher plant. Am.J. Bot. 85:88–91.OPLINGER, C. S. 1967. Food habits and feeding activity of recently transformedand adult Hyla crucifer crucifer Wied. Herpetologica 23:209–217.SCHNELL, D. E. 2002. Carnivorous Plants of the United States and Canada.Timber Press, Portland, Oregon.THE SENTINEL. 2007. Plants Hungry for Meat, May 01. Danville, California.WRIGHT, A. H., AND A. A. WRIGHT. 1949. Handbook of Frogs and Toads ofthe United States and Canada, 3 rd ed. Comstock Publishing Associates,Ithaca, New York. 640 pp.<strong>Herpetological</strong> <strong>Review</strong> 39(2), 2008 155
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FIG. 1. Oscillogram, spectrogram, a
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FIG. 1. Adult Physalaemus cuvieri r
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Répteis, Instituto Nacional de Pes
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discovered 145 live hatchlings and
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GRAPTEMYS GIBBONSI (Pascagoula Map
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College, and the Joseph Moore Museu
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FIG. 1. Common Ground Lizard (Ameiv
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havior unavailable elsewhere. Here
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15% of predator mass, is typical fo
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side the third burrow and began a f
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We thank Arlington James and the st
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mm) S. viridicornis in its mouth in
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NECTURUS MACULOSUS (Common Mudpuppy
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LITHOBATES CATESBEIANUS (American B
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Research and Collections Center, 13
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BRONCHOCELA VIETNAMENSIS (Vietnam L
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Oficina Regional Guaymas, Guaymas,
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MICRURUS TENER (Texas Coralsnake).
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declining in this recently discover
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80.7372°W). 02 November 2005. Stev
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this effort, 7% of the 10 × 10 km
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the knowledge of the group. The aut
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which is listed under “Rhodin, A.
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noting that Sphenomorphus bignelli
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ISSN 0018-084XThe Official News-Jou