Pennsylvania Land Snails


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ANDERSON, R.C. and A.K. PRESTWOOD. 1981. Parelaphostrongylosis. In Diseases and Parasites of White-tailed Deer, W.R. Davidson, Ed. Tall Timbers Research Station, Tallahassee, FL. pp. 274-308.


ATKINSON, J.W. 1998. Food Manipulation and Transport by a Carnivorous Land Snail, Haplotrema concavum. Invert. Biol. 117(2):109-113.


___. 2003. Foraging strategy switch in detour behavior of the land snail Anguispira alternata (Say). Invertebrate Biology 122 (4):326-333.


BARKER, G.M., Ed. 2004. Natural Enemies of Terrestrial Molluscs. CABI Publishing, Oxford, UK. 644pp.


BAUR, B, and BAUR, A.1993. Climatic warming due to thermal radiation from an urban area as a possible cause for the local extinction of a land snail. Journal of Applied Ecology 30: 333-340.


BIRD, J. and J.M. GARVON. 2005. Attraction of the land snail Anguispira alternata to fresh faeces of white-tailed deer: implications in the transmission of Parelaphostrongylus tenuis. Canadian Journal of Zoology 83(2):358-362.


BLINN, W.C. 1963. Ecology of the land snails Mesodon thyroidus and Allogona profunda. Ecology 44(3):498-505.


BOBKOVA, M.V., O.S. TARTAKOVSKAYA, S.L. BORISSENKO, V.V. ZHUKOV and V.B. MEYER-ROCHOW. 2004. Restoration of morphological and functional integrity in the regenerating eye of the giant African land snail Achatina fulica. Acta Zoologica 85:1-14.


BOETTCHER, S.E. and P.J. KALISZ. 1990. Single-tree influence on soil properties in the mountains of eastern Kentucky. Ecology 71(4):1365-1372.


BROOKS, S.T. 1931. A list of the land-snails of Pennsylvania with a summary of their distributions. Annals of the Carnegie Museum 22 (3-4): 313-331.


BURCH, J.B. 1955. 1962. How To Know the Eastern Land Snails. Wm. C. Brown, Dubuque. 214 pp.


___. 1955. Some ecological factors of the soil affecting the distribution and abundance of land snails in eastern Virginia. Nautilus 69(2):26-29.


___. 1990. Thomas Say’s glossary for conchology. Walkerana 4:279-306.


CONEY, C.C., W.A. TARPLEY, J.C. WARDEN and J.W. NAGEL. 1982. "Ecological studies of land snails in the Hiwassee River Basin of Tennessee," U.S.A. Malacological Review 15: 69-106.


DALLINGER, R. and W. WIESER. 1984. Patterns of accumulation, distribution and liberation of Zn, Cu, Cd and Pb in different organs of the land snail Helix pomatia L. Comparative Biochemistry and Physiology - Part C: Toxicology and Pharmacology 79(1):117-124.


DAVISON, A., C.M. WADE, M. CHRISTOPHER, P.B. MORDAN and S. CHIBA. 2005. Sex and darts in slugs and snails (Mollusca: Gastropoda: Stylommatophora). Journal of Zoology, London 267: 329-338.


DeCALESTA, D.S. 1994. Effect of white-tailed deer on songbirds within managed forests in Pennsylvania. Journal of Wildlife Management 58(4):711-718.


DIGWEED, S.C. 1993. Selection of terrestrial gastropod prey by Cychrine and Pterostichine ground beetles (Coleoptera: Carabidae). The Canadian Entomologist 125: 463-472.


DIMITRIADIS, V.K. 2001. Chapter 5: Structure and Function of the Digestive System of Stylommatophora. Pp 237-257 in The Biology of Terrestrial Molluscs, Ed. G.M. Barker, CABI, New York, USA. 558 pp.


EISNER, T. and E.O. WILSON. 1970. Defensive liquid discharge in Florida tree snails (Liguus fasciatus). The Nautilus 84(1):14-15.


ELWELL, A.S. and M.J. ULMER. 1971. Notes on the biology of Anguispira alternata (Stylommatophora: Endodontidae). Malacologia 11(1):199-215.


EMBERTON, K.C. 1988. The genitalic, allozymic and conchological evolution of the eastern North American Triodopsinae (Gastropoda: Pulmonata: Polygyridae). Malacologia 28(1-2): 159-273.


___. 1991. The genitalic, allozymic and conchological evolution of the tribe Mesodontini (Pulmonata: Stylommatophora: Polygyridae). Malacologia 33(1-2):71-178.


___. 1995. When shells do not tell: 145 million years of evolution in North America's polygyrid land snails, with a revision and conservation priorities. Malacologia 37(1):69-110.


FAIRBANKS, H.L. 1986. The taxonomic status of Philomycus togatus (Pulmonata:Philomycidae): a morphological and electrophoretic comparison with Philomycus carolinianus. Malacologia 27(2):271-280.


GLEICH, J.G., F.F. GILBERT and N.P. KUTSCHA. 1977. Nematodes in terrestrial gastropods from central Maine. Journal of Wildlife Diseases 13:43-46.


GOMEZ, B.J. 2001. Chapter 8: Structure and Functioning of the Reproductive System. Pp 307-330 in The Biology of Terrestrial Molluscs, G.M. Barker, Ed., CABI, New York. 558 pp.


GRAVELAND, J.R. 1996. Avian eggshell formation in calcium-rich and calcium-poor habitats: importance of snail shells and anthropogenic calcium sources. Canadian Journal of Zoology 74:1035-1044.


GRAVELAND, J., R. van der WAL, J.H. van BALEN and A.J. van NOORDWIJK. 1994. Poor reproduction in forest passerines from decline of snail abundance on acidified soils. Nature 368: 446-448.


GRIME, J.P. and G.M. BLYTHE. 1969. An investigation of the relationship between snails and vegetation at the Winnats Pass. Journal of Ecology 57: 45-66.


HAMBURG, S.P. R.D. YANAI, M.A. ARTHUR, J.D. BLUM and T.G. SICCAMA. 2003. Biotic control of calcium cycling in northern hardwood forests: acid rain and aging forests. Ecosystems 6:399-406.


HAMES, R.S., K.V. ROSENBERG, J.D. LOWE, S.E. BARKER and A.A. DHONDT. 2002. Adverse effects of acid rain on the distribution of the wood thrush Hylocichla mustelina in North America. Proceedings of the National Academy of Science 99(17):11235-11240.


HANLEY, M.E., M. FENNER, and P.J. EDWARDS. 1995. An experimental field study of the effects of mollusc grazing on seedling recruitment and survival in grassland. Journal of Ecology 83:621-627.


HELLER, J. 2001. Chapter 12: Life History Strategies. Pp 413-445 in The Biology of Terrestrial Molluscs, G.M. Barker, Ed., CABI, New York. 558 pp.


HOBBS, H.H. III. 1995. First record of the troglophilic terrestrial snail Carychium exile Lea from Indiana caves (Gastropoda: Stylommatophora: Carychiidae). National Speleological Society Bulletin 56:104-105.


HOTOPP, K.P. 2002. Land snails and soil calcium in central Appalachian Mountain forest. Southeastern Naturalist 1(1):27-44.


HUBRICHT, L. 1985. The distributions of the native land mollusks of the eastern United States. Fieldiana: Zoology, New Series, No. 24: 1-191.


___. 1962. New species of Helicodiscus from the eastern United States. Nautilus 75(3):102-108.


___. 1951. The Limacidae and Philomycidae of Pittsylvania County, Virginia. The Nautilus 65(1):20-22.


INGRAM, W.M. 1942. Snail associates of Blarina brevicauda talpoides. Journal of Mammalogy, 23: 255-258.


JOHANNESSEN, l.E. and T. SOLHØY. 2001. Effects of experimentally increased calcium levels in the litter on terrestrial snail populations. Pedobiologia 45(3):234-242.


JORDAENS, K., T. BACKELJAU, P. ODINA, H. REISE and R. VERHAGEN. 1998. Allozyme homozygosity and phally polymorphism in the land snail Zonitoides nitidus (Gastropoda, Pulmonata). Journal of Zoology 246:95-104.

    KADO, Y. 1960. Studies on shell formation in molluscs. Journal of Science of Hiroshima University 19:163-210.  

KERNEY, M.P. and R.A.D. CAMERON. 1979. A Field Guide to the Land Snails of Britain and North-west Europe. Collins, London. 288 pp.


KOENE, J. M. and R. CHASE. 1998. Changes in the reproductive system of the snail Helix aspersa caused by mucus from the love dart. Journal of Experimental Biology 201: 2313-2319.

    LEADY, B.S. and J.F. GOTTGENS. 2001. Mercury accumulation in sediment cores and along food chains in two regions of the Brazilian Pantanal. Wetlands Ecology and Management 9:349-361.  

LLOYD, M. 1967. Numerical observations on movements of animals between beech litter and fallen branches. Journal of Animal Ecology 36:1-30.


MACKENSTEDT, U. and K. MÄRKEL. 2001. Chapter 4: Radular Structure and Function, Pp 213-236 in The Biology of Terrestrial Molluscs, Ed. G.M. Barker, CABI, New York, USA. 558 pp.

    MacMILLAN, G.K. 1950. The occurrence of Hendersonia occulta (Say) in Pennsylvania. Proceeedings of the Pennsylvania Academy of Science 24:113-115.  
    MARTIN, A.C., H.S. ZIM and A.L. NELSON. 1951. American Wildlife and Plants: A Guide to Wildlife Food Habits. Dover, New York. 500 pp.  
    MASON, C.F. 1970. Snail populations, beech litter production, and the role of snails in litter decomposition. Oecologia 5:215-239.  
    MAZE, R.J. and C. JOHNSTONE. 1984. Gastropod intermediate hosts of the meningeal worm Parelaphostrongylus tenuis in Pennsylvania: observations on their ecology. Canadian Journal of Zoology 64:185-188.  
    McCRACKEN, G.F. and P.F. BRUSSARD. 1980. Self-fertilization in the white-lipped snail Triodopsis albolabris. Biological Journal of the Linnaean Society 14:429-434.  
    NEKOLA, J.C. 1999. Terrestrial gastropod richness of carbonate cliff and associated habitats in the Great Lakes Region of North America. Malacologia 41(1):231-252.  

___ and M. BARTHEL. 2002. Morphometric analysis of Carychium exile and Carychium exiguum in the Great Lakes region of North America. Journal of Conchology 37(5):515.


PARKARINEN, E. 1994. The importance of mucus as a defence against carabid beetles by the slugs Arion fasciatus and Deroceras reticulatum. Journal of Molluscan Studies 60:149-155.


PARKER, G.R. 1966. Moose disease in Nova Scotia; gastropod-nematode relationship. Unpublished MS thesis, Acadia University, Wolfville, N.S. 126 pp.


PEARCE, T.A. and A. GAERTNER. 1996. Optimal foraging and mucus-trail following in the snail-eating snail Haplotrema concavum (Gastropoda: Pulmonata). Malacological Review 29:85-99.


PENNSYLVANIA DEPARTMENT of CONSERVATION and NATURAL RESOURCES. 2005. Forests: Lack of Regeneration. ( Online report accessed 12/26/05.


PILSBRY, H.A. 1940. Land Mollusca of North America (North of Mexico), Vol. 1, Part 2. Monograph No. 3, Academy of Natural Sciences of Philadelphia. Philadelphia, PA. 994 pp.


___. 1946-1948. Land Mollusca of North America. Monograph of the Academy of Natural Sciences of Philadelphia, 3(2). 1102 pp.


REGOLI, F., S. GORBI, D. FATTORINI, S. TEDESCO, A. NOTTI, N. MACHELLA, R. BOCCHETTI, M. BENEDETTI and F. PIVA. 2006. Use of the land snail Helix aspersa as a sentinel organism for monitoring ecotoxicologic effects of urban pollution: an integrated approach. Environmental Health Perspectives 114(1).


RIMMER, C.C., K. P. McFARLAND, D. C. EVERS, E. K. MILLER, Y. AUBRY, D. BUSBY, and R. J. TAYLOR. 2005. Mercury levels in Bicknell's thrush and other insectivorous passerine birds in montane forests of the northeastern United States and Canada. Ecotoxicology 14:223-240.


ROTH, B. 1991. A phylogenetic analysis and revised classification of the North American Haplotrematidae (Gastropoda: Pulmonata). American Malacological Bulletin, 8: 155-163.


SCHEIFLER, R., C, SCHWARTZ, G. ECHEVARRIA, A. de VAUFLEURY, P.-M. BADOT and J.-L. MORE. 2003a. "Nonavailable" soil cadmium is bioavailable to snails: evidence from isotopic dilution experiments. Environmental Science and Technology 37(1):81-86.


SCHILTHUIZEN, M. 2005. The darting game in snails and slugs. Trends in Ecology and Evolution 20(11): 581-584.


SOLEM, A. 1972. Microarmature and barriers in the aperture of land snails. The Veliger 15(2):81-87.


SPEISER, B. 2001. Chapter 6: Food and feeding behavior, pp. 259- 288 in The Biology of Terrestrial Molluscs, Ed. G.M. Barker, CABI, New York, USA. 558 pp.


STRÖM, L. 2004. Long-term effects of riparian clear-cutting —richer land snail communities in regenerating forests. Unpublished MS thesis, Umeå University, Sweden. 23 pp.


SUOMINEN, O. 1999. Impact of cervid browsing and grazing on the terrestrial gastropod fauna in the boreal forests of Fennoscandia. Ecography 22:651-658.


SUOMINEN, O., K. DANELL and R. BERGSTRÖM. 1999. Moose, trees and ground-living invertebrates: indirect interactions in Swedish pine forests. Oikos 84:215-226.


SUVOROV, A.N. 1993. Some aspects of functional morphology of the aperture in the Pupillina suborder (Gastropoda Pulmonata). Ruthenica 3:141–152.


___. 1999. Functional interrelations between aperture structures and soft organs in lower Geophila: 1. Pupillina, Oleacinina. Russian Journal of Zoology 3:5–15.


SYMONDSON, W.O.C. 2004. Chapter 2: Coleoptera (Carabidae, Staphylinidae, Lampyridae, Drilidae and Silphidae) as predators of terrestrial gastropods, pp. 37-84 in Natural Enemies of Terrestrial Molluscs. CABI Publishing, Oxford, UK. 644pp.


UPSHALL, S.M., M.D.B. BURT and T.G. DILWORTH. 1985. Parelaphostrongylus tenuis in New Brunswick: the parasite in terrestrial gastropods. Journal of Wildlife Diseases 22(4):582-586.


VALOVIRTA, I. 1968. Land mollusks in relation to acidity on hyperite hills in central Finland. Annales Zoologici Fennici 5:245-253.


VESTERDAHL, L. and K. RAULAND-RASMUSSEN. 1998. Forest floor chemistry under seven tree species along a soil fertility gradient. Canadian Journal of Forest Research 28:1636-1647.


WARDLE, D.A., G.M. BARKER, G.W. YEATES, K.I. BONNER and A. GHANI. 2001. Introduced browsing mammals in natural New Zealand forests: aboveground and belowground consequences. Ecological Monographs 71:587-614.


WÄREBORN, I. 1992. Changes in the land mollusc fauna and soil chemistry in an inland district in southern Sweden. Ecography 15: 62-69.


WURTZ, C.B. 1940. New state and county records of land-snails in Pennsylvania. Nautilus 53(3): 84-88.


YANAI, R.D., T.G. SICCAMA, M.A. ARTHUR, C.A. FEDERER, and A.J. FRIEDLAND. 1999. Accumulation and depletion of base cations in forest floors in the Northeastern United States. Ecology 80(8):2771-2787.

Content for this website is written by Ken Hotopp, Principal of Appalachian Conservation Biology, and edited by Tim Pearce, Head of Carnegie Museum of Natural History’s Section of Mollusks. The website was developed by Cathy Klingler, Carnegie Museum of Natural History's Web Site Manager. Snail images are by Ron Lutz II, Larry Watrous, and Ed Thompson.

Cite as:
Hotopp, K.P., and T.A. Pearce. 2005. Land Snails of Pennsylvania. Carnegie Museum of Natural History, Pittsburgh, PA, USA.


PA DCNRDevelopment of this site was supported by the generous contributions of Pennsylvanians to the Wild Resource Conservation Fund.