Using landscape ecology to understand and manage freshwater mussel populations

Journal of the North American Benthological Society

Volumn 27, Issue 2, 2008, Pages 424-439

  Newton, Teresa J ^a   Woolnough, Daelyn A ^b   Strayer, David L ^c  

^a NATIONAL WILDLIFE HEALTH CENTER   (United States)

^b TRENT UNIVERSITY   (Canada)

^c CARY INSTITUTE OF ECOSYSTEM STUDIES   (United States)

Author keywords

Connectivity; Freshwater mussels; Habitat; Landscape ecology; Patches; Spatial scales; Watershed processes

Indexed keywords

BIVALVE; CONNECTIVITY; HABITAT FRAGMENTATION; HUMAN ACTIVITY; HYDRAULICS; LANDSCAPE ECOLOGY; PATCH DYNAMICS; WATERSHED;

EID: 47049102092     PISSN: 08873593     EISSN: None     Source Type: Journal    
DOI: 10.1899/07-076.1     Document Type: Review

References (119)

1. ADDICOTT, J. F., J. M. AHO, M. F. ANTOLIN, D. K. PADILLA, J. S. RICHARDSON, AND D. A. SOLUK. 1987. Ecological neighborhoods: scaling environmental patterns. Oikos 49:340-346.
2. AGNEW, L. J., S. LYON, P. GÉRARD- MARCHANT, V. B. COLLINS, A. J. LEMBO, T. S. STEENHUIS, AND M. T. WALTER. 2006. Identifying hydrologically sensitive areas: bridging the gap between science and application. Journal of Environmental Management 78:63-76.
3. ALLAN, J. D. 2004. Landscapes and riverscapes: the influence of land use on stream ecosystems. Annual Review of Ecology and Systematics 35:257-284.
4. ANDERSON, D. R., K. P. BURNHAM, W. R. GOULD, AND S. CHERRY. 2001. Concerns about finding effects that are actually spurious. Wildlife Society Bulletin 29:311-316.
5. ARBUCKLE, K. E., AND J. A. DOWNING. 2002. Freshwater mussel abundance and species richness: GIS relationships with watershed land use and geology. Canadian Journal of Fisheries and Aquatic Sciences 59:310-316.
6. AUGSPURGER, T., A. E. KELLER, M. C. BLACK, W. G. COPE, AND F. J. DWYER. 2003. Water quality guidance for protection of freshwater mussels (Unionidae) from ammonia exposure. Environmental Toxicology and Chemistry 22:2569-2575.
7. BAUER, G. 1988. Threats to the freshwater pearl mussel Margaritifera margaritifera L. in central Europe. Biological Conservation 45:239-253.
8. BERTNESS, M. D., AND E. GROSHOLZ. 1985. Population dynamics of the ribbed mussel, Geukensia demissa: the costs and benefits of an aggregated distribution. Oecologia (Berlin) 67:192-204.
9. BRAINWOOD, M., S. BURGIN, AND M. BYRNE. 2006. Is the decline of freshwater mussel populations in a regulated coastal river in southeastern Australia linked with human modification of habitat? Aquatic Conservation-Marine and Freshwater Ecosystems 16:501-516.
10. BRIERLEY, G. J., T. COHEN, K. FRYIRS, AND A. BROOKS. 1999. Post-European changes to the fluvial geomorphology of Bega catchment, Australia: implications for river ecology. Freshwater Biology 41:839-848.
11. BRIM BOX, J., R. M. DORAZIO, AND W. D. LIDDELL. 2002. Relationships between streambed substrate characteristics and freshwater mussels (Bivalvia:Unionidae) in coastal plain streams. Journal of the North American Benthological Society 21:253-260.
12. CHURCH, M. 2002. Geomorphic thresholds in riverine landscapes. Freshwater Biology 47:541-557.
13. CLARKE, R. T., J. A. THOMAS, G. W. ELMES, AND M. E. HOCHBERG. 1997. The effects of spatial patterns in habitat quality on community dynamics within a site. Proceedings of the Royal Society of London: Series B-Biological Sciences 264:347-354.
14. COKER, R. E., A. F. SHIRA, H. W. CLARK, AND A. D. HOWARD. 1921. Natural history and propagation of fresh-water mussels. Bulletin of the Bureau of Fisheries 37:77-181.
15. COOPER, S. D., L. BARMUTA, O. SARNELLE, K. KRATZ, AND S. DIEHL. 1997. Quantifying spatial heterogeneity in streams. Journal of the North American Benthological Society 16:174-188.
16. DENNIS, R. L. H., AND H. T. EALES. 1997. Patch occupancy in Coenonympha tullia (Müller, 1764) (Lepidoptera: Satyrinae): habitat quality matters as much as patch size and occupancy. Journal of Insect Conservation 1:167-176.
17. DI MAIO, J., AND L. D. CORKUM. 1995. Relationship between the spatial distribution of fresh-water mussels (Bivalvia, Unionidae) and the hydrological variability of rivers. Canadian Journal of Zoology 73:663-671.
18. ECKBLAD, J. 1999. Evaluation of unionid mussel colonization of dredge cuts and dredged material placement sites in Pool 11-22 of the upper Mississippi River. Final report (DACW 25-95-D-006) to the US Army Corps of Engineers, Rock Island, Illinois. (Available from: US Army Engineer District, Clock Tower Building, P.O. Box 2004, Rock Island, Illinois 61204 USA.)
19. ELLIS, M. M. 1936. Erosion silt as a factor in aquatic environments. Ecology 17:29-42.
20. ENGEN, S., R. LANDE, AND B.-E. SAETHER. 2002. Migration and spatiotemporal variation in population dynamics in a heterogeneous environment. Ecology 83:570-579.
21. FAGAN, W. F. 2002. Connectivity, fragmentation, and extinction risk in dendritic metapopulations. Ecology 83:3243-3249.
22. FAHRIG, L. 2003. Effects of habitat fragmentation on biodiversity. Annual Review of Ecology, Evolution, and Systematics 34:487-515.
23. FAHRIG, L., AND G. MERRIAM. 1985. Habitat patch connectivity and population survival. Ecology 66:1762-1768.
24. FAHRIG, L., AND W. K. NUTTLE. 2005. Population ecology in spatially heterogeneous environments. Pages 95-118 in G. M. Lovett, C. G. Jones, M. G. Turner, and K. C. Weathers (editors). Ecosystem function in heterogeneous landscapes. Springer, New York.
25. FLEMING, W. J., T. P. AUGSPURGER, AND J. A. ALDERMAN. 1995. Fresh-water mussel die-off attributed to anticholinesterase poisoning. Environmental Toxicology and Chemistry 14:877-879.
26. FORMAN, R. T. T., AND M. GODRON. 1981. Patches and structural components for a landscape ecology. BioScience 31:733-740.
27. FORTIN, M.-J., AND M. DALE. 2005. Spatial analysis: a guide for ecologists. Cambridge University Press, Cambridge, UK.
28. GAGNON, P., W. MICHENER, M. FREEMAN, AND J. BRIM BOX. 2006. Unionid habitat and assemblage composition in coastal plain tributaries of Hint River (Georgia). Southeastern Naturalist 5:31-52.
29. GORDON, N. D., T. A. MCMAHON, AND B. L. FINLAYSON. 1992. Stream hydrology: an introduction for ecologists. John Wiley and Sons, Chichester, UK.
30. GRIFFITH, D. R. 2003. Spatial autocorrelation and spatial filtering. Springer, New York.
31. HAAG, W. R., D. J. BERG, D. W. GARTON, AND J. L. FARRIS. 1993. Reduced survival and fitness in native bivalves in response to fouling by the introduced zebra mussel (Dreissena polymorpha) in western Lake Erie. Canadian Journal of Fisheries and Aquatic Sciences 50:13-19.
32. HAAG, W. R., AND M. L. WARREN. 1998. Role of ecological factors and reproductive strategies in structuring freshwater mussel communities. Canadian Journal of Fisheries and Aquatic Sciences 55:297-306.
33. HADDAD, N. M., D. R. BOWNE, A. CUNNINGHAM, B. J. DANIELSON, D. J. LEVEY, S. SARGENT, AND T. SPIRA. 2003. Corridor use by diverse taxa. Ecology 84:609-615.
34. HANSKI, I. 1998. Metapopulation dynamics. Nature 396:41-49.
35. HANSKI, I. 1999. Habitat connectivity, habitat continuity and metapopulations in dynamic landscapes. Oikos 87:209-219.
36. HANSKI, I. 2002. Metapopulation ecology. Oxford University Press, New York.
37. HANSKI, I., AND O. E. GAGGIOTTI. 2004. Ecology, genetics, and evolution of metapopulations. Elsevier Academic Press, New York.
38. HARDISON, B. S., AND J. B. LAYZER. 2001. Relations between complex hydraulics and the localized distribution of mussels in three regulated rivers. Regulated Rivers: Research and Management 17:77-84.
39. HARPER, D. M., K. L. KEMP, B. VOGEL, AND M. D. NEWSON. 2000. Towards the assessment of 'ecological integrity' in running waters of the United Kingdom. Hydrobiologia 422:133-142.
40. HARRISON, S., AND E. BRUNA. 1999. Habitat fragmentation and large-scale conservation: what do we know for sure? Ecography 22:225-232.
41. HARTFIELD, P. 1993. Headcuts and their effect on freshwater mussels. Pages 131-141 in K. S. Cummings, A. C. Buchanan, and L. M. Koch (editors). Conservation and management of freshwater mussels. Upper Mississippi River Conservation Committee, Rock Island, Illinois.
42. HASTIE, L. C., S. L. COOKSLEY, F. SCOUGALL, M. R. YOUNG, P. J. BOON, AND M. J. GAYWOOD. 2004. Application of extensive survey techniques to describe freshwater pearl mussel distribution and macrohabitat in the River Spey, Scotland. River Research and Applications 20:1001-1013.
43. HASTIE, L. C., P. J. COSGROVE, N. ELLIS, AND M. J. GAYWOOD. 2003. The threat of climate change to freshwater pearl mussel populations. Ambio 32:40-46.
44. HEINRICHER, J. R., AND J. B. LAYZER. 1999. Reproduction by individuals of a nonreproducing population of Megalonaias nervosa (Mollusca: Unionidae) following translocation. American Midland Naturalist 141:140-148.
45. HOULAHAN, J. E., AND C. S. FINDLAY. 2004. Estimating the "critical" distance at which adjacent land use degrades wetland water and sediment quality. Landscape Ecology 19:677-690.
46. HOWARD, J. K., AND K. M. CUFFEY. 2003. Freshwater mussels in a California North Coast Range river: occurrence, distribution, and controls. Journal of the North American Benthological Society 22:63-77.
47. HUGHES, J. M. 2007. Constraints on recovery: using molecular methods to study connectivity of aquatic biota in rivers and streams. Freshwater Biology 52:616-631.
48. HUGHES, R. M., L. W. WANG, AND P. W. SEELBACH. 2006. Landscape influences on stream habitats and biological assemblages. American Fisheries Society Symposium 48. American Fisheries Society, Bethesda, Maryland.
49. IMLAY, M. J. 1972. Greater adaptability of freshwater mussels to natural rather than to artificial displacement. Nautilus 86:76-79.
50. JAGER, H. I., J. A. CHANDLER, K. B. LEPLA, AND W. VAN WINKLE. 2001. A theoretical study of river fragmentation by dams and its effect on white sturgeon populations. Environmental Biology of Fishes 60:347-361.
51. JOHNSON, P. D., AND K. M. BROWN. 2000. The importance of microhabitat factors and habitat stability to the threatened Louisiana pearl shell, Margaritifera hembeli (Conrad). Canadian Journal of Zoology 78:271-277.
52. JOHNSON, T. E., J. N. MCNAIR, P. SRIVASTAVA, AND D. D. HART. 2007. Stream ecosystem responses to spatially variable land cover: an empirically based model for developing riparian restoration strategies. Freshwater Biology 52:680-695.
53. KELLY, M., AND J. RHYMER. 2005. Population genetic structure of a rare unionid (Lampsilis cariosa) in a recently glaciated landscape. Conservation Genetics 6:789-802.
54. KELNER, D. E., AND B. E. SIETMAN. 2000. Relic populations of the ebony shell, Fusconaia ebena (Bivalvia: Unionidae) in the upper Mississippi River drainage. Journal of Freshwater Ecology 15:371-377.
55. KEMP, J. L., D. M. HARPER, AND G. A. CROSA. 2000. The habitat-scale ecohydraulics of rivers. Ecological Engineering 16:17-29.
56. KILLEEN, I., D. ALDRIDGE, AND G. OLIVER. 2004. Freshwater bivalves of Britain and Ireland. National Museum of Wales Occasional Publication 82:1-114.
57. LAYZER, J. B., AND L. M. MADISON. 1995. Microhabitat use by freshwater mussels and recommendations for determining their instream flow needs. Regulated Rivers: Research and Management 10:329-345.
58. LEE, H.-L., D. DEANGELIS, AND H.-L. KOH. 1998. Modeling spatial distribution of the unionid mussels and the core-satellite hypothesis. Water Science and Technology 38:73-79.
59. LEOPOLD, L. B., M. G. WOLMAN, AND J. P. MILLER. 1964. Fluvial processes in geomorphology. W. H. Freeman, San Francisco, California.
60. LEVIN, S. A., J. H. STEELE, AND T. M. POWELL. 1993. Patch dynamics. Springer, New York.
61. LYONS, J., S. W. TRIMBLE, AND L. K. PAINE. 2000. Grass versus trees: managing riparian areas to benefit streams of central North America. Journal of the American Water Resources Association 36:919-930.
62. MADON, S. P., D. W. SCHNEIDER, J. A. STOECKEL, AND R. E. SPARKS. 1998. Effects of inorganic sediment and food concentrations on energetic processes of the zebra mussel, Dreissena polymorpha: implications for growth in turbid rivers. Canadian Journal of Fisheries and Aquatic Sciences 55:401-413.
63. MALANSON, G. P. 2003. Dispersal across continuous and binary landscapes. Ecological Modelling 169:17-24.
64. MALANSON, G. P., AND B. E. CRAMER. 1999. Landscape heterogeneity, connectivity, and critical landscapes for conservation. Diversity and Distributions 5:27-39.
65. MALMQVIST, B., AND S. RUNDLE. 2002. Threats to the running water ecosystems of the world. Environmental Conservation 29:134-153.
66. MCCULLOUGH, D. 1996. Metapopulations and wildlife conservation management. Island Press, Washington, DC.
67. MCRAE, S. E., J. D. ALLAN, AND J. B. BURCH. 2004. Reach- and catchment-scale determinants of the distribution of freshwater mussels (Bivalvia: Unionidae) in southeastern Michigan, USA. Freshwater Biology 49:127-142.
68. METCALFE-SMITH, J. L., S. K. STATON, G. L. MACKIE, AND N. M. LANE. 1998. Changes in the biodiversity of freshwater mussels in the Canadian waters of the lower Great Lakes drainage basin over the past 140 years. Journal of Great Lakes Research 24:845-858.
69. MOLES, K. R., AND J. B. LAYZER. 2008. Reproductive ecology of Actinonaias ligamentina (Bivalvia:Unionidae) in a regulated river. Journal of the North American Benthological Society 27:212-222.
70. MORALES, Y., L. J. WEBER, A. E. MYNETT, AND T. J. NEWTON. 2006. Effects of substrate and hydrodynamic conditions on the formation of mussel beds in a large river. Journal of the North American Benthological Society 25:664-676.
71. MORRIS, T. J., AND L. D. CORKUM. 1996. Assemblage structure of freshwater mussels (Bivalvia:Unionidae) in rivers with grassy and forested riparian zones. Journal of the North American Benthological Society 15:576-586.
72. MORRIS, T. J., AND L. D. CORKUM. 1999. Unionid growth patterns in rivers of differing riparian vegetation. Freshwater Biology 42:59-68.
73. MOULTON, C. A., W. J. FLEMING, AND C. E. PURNELL. 1996. Effects of two cholinesterase-inhibiting pesticides on freshwater mussels. Environmental Toxicology and Chemistry 15:131-137.
74. NAIMAN, R. J., H. DÉCAMPS, AND M. E. MCCLAIN. 2005. Riparia: ecology, conservation, and management of streamside communities. Elsevier, Amsterdam, Netherlands.
75. NAIMO, T. J. 1995. A review of the effects of heavy metals on freshwater mussels. Ecotoxicology 4:341-362.
76. NAKANO, S., F. KITANO, AND K. MAEKAWA. 1996. Potential fragmentation and loss of thermal habitats for charrs in the Japanese archipelago due to climatic warming. Freshwater Biology 36:711-722.
77. NICHOLS, S. J., AND D. GARLING. 2000. Food-web dynamics and trophic-level interactions in a multispecies community of freshwater unionids. Canadian Journal of Zoology 78:871-882.
78. ODUM, E. P. 1971. Fundamentals of ecology. W. B. Saunders, Philadelphia, Pennsylvania.
79. O'NEILL, R. V., D. L. DEANGELIS, J. B. WAIDE, AND T. F. H. ALLEN. 1986. A hierarchical concept of ecosystems. Princeton University Press, Princeton, New Jersey.
80. ORTMANN, A. E. 1909. The destruction of the fresh-water fauna in western Pennsylvania. Proceedings of the American Philosophical Society 48:90-110.
81. ORTMANN, A. E. 1919. A monograph of the naiades of Pennsylvania. Part III: systematic account of the genera and species. Memoirs of the Carnegie Museum 8:1-384.
82. PALMER, M. A., C. M. SWAN, K. NELSON, P. SILVER, AND R. ALVESTAD. 2000. Streambed landscapes: evidence that stream invertebrates respond to the type and spatial arrangement of patches. Landscape Ecology 15:563-576.
83. PEACOCK, E., W. R. HAAG, AND M. L. WARREN. 2005. Prehistoric decline in freshwater mussels coincident with the advent of maize agriculture. Conservation Biology 19:547-551.
84. PEARSON, S. M., M. G. TURNER, R. H. GARDNER, AND R. V. O'NEILL. 1996. An organism-based perspective of habitat fragmentation. Pages 77-95 in R. C. Szaro and D. W. Johnston (editors). Biodiversity in managed landscapes: theory and practice. Oxford University Press, New York.
85. POFF, N. L., AND J. V. WARD. 1990. Physical habitat template of lotic systems: recovery in the context of spatiotemporal heterogeneity. Environmental Management 14:629-645.
86. POOLE, K. E., AND J. A. DOWNING. 2004. Relationship of declining mussel biodiversity to stream-reach and watershed characteristics in an agricultural landscape. Journal of the North American Benthological Society 23:114-125.
87. SMITH, D. G. 1985. Recent range expansion of the freshwater mussel Anodonta implicata and its relationship to clupeid fish restoration in the Connecticut River system. Freshwater Invertebrate Biology 4:105-108.
88. SPARKS, B. L., AND D. L. STRAYER. 1998. Effects of low dissolved oxygen on juvenile Elliptio complanata (Bivalvia:Unionidae). Journal of the North American Benthological Society 17:129-134.
89. STRAYER, D. L. 1983. The effects of surface geology and stream size on freshwater mussel distribution in southeastern Michigan, USA. Freshwater Biology 13:253-264.
90. STRAYER, D. L. 1999. Use of flow refuges by unionid mussels in rivers. Journal of the North American Benthological Society 18:468-476.
91. STRAYER, D. L. 2008. Freshwater mussel ecology: a multifactor approach to distribution and abundance. University of California Press, Berkeley, California.
92. STRAYER, D. L., R. E. BEIGHLEY, L. C. THOMPSON, S. BROOKS, C. NILSSON, G. PINAY, AND R. J. NAIMAN. 2003. Effects of landcover change on stream ecosystems: roles of empirical models and scaling issues. Ecosystems 6:407-423.
93. STRAYER, D. L., J. A. DOWNING, W. R. HAAG, T. L. KING, J. B. LAYZER, T. J. NEWTON, AND S. J. NICHOLS. 2004. Changing perspectives on pearly mussels, North America's most imperiled animals. BioScience 54:429-439.
94. STRAYER, D. L., AND J. RALLEY. 1993. Microhabitat use by an assemblage of stream-dwelling unionaceans (Bivalvia), including two rare species of Alasmidonta. Journal of the North American Benthological Society 12:247-258.
95. SVANE, I., AND M. OMPI. 1993. Patch dynamics in beds of the blue mussel Mytilus edulis L.: effects of site, patch size, and position within a patch. Ophelia 37:187-202.
96. TAYLOR, P. D., L. FAHRIG, K. HENEIN, AND G. MERRIAM. 1993. Connectivity is a vital element of landscape structure. Oikos 68:571-573.
97. THOMPSON, J. N. 1978. Within-patch structure and dynamics in Pastinaca sativa and resource availability to a specialized herbivore. Ecology 59:443-448.
98. TILMAN, D., R. M. MAY, C. L. LEHMAN, AND M. A. NOWAK. 1994. Habitat destruction and the extinction debt. Nature 371:65-66.
99. TRIMBLE, S. W. 1981. Changes in sediment storage in the Coon Creek basin, Driftless Area, Wisconsin, 1853 to 1975. Science 214:181-183.
100. TURNER, M. G., R. H. GARDNER, AND R. V. O'NEILL. 2001. Landscape ecology in theory and practice. Springer-Verlag, New York.
101. VAN DER SCHALIE, H. 1938. Contributing factors in the depletion of naiades in the eastern United States. Basteria 3:51-57.
102. VAUGHN, C. C., K. B. GIDO, AND D. E. SPOONER. 2004. Ecosystem processes performed by unionid mussels in stream mesocosms: species roles and effects of abundance. Hydrobiologia 527:35-47.
103. VAUGHN, C. C., AND C. M. TAYLOR. 2000. Macroecology of a host-parasite relationship. Ecography 23:11-20.
104. WALLER, D. L., S. GUTREUTER, AND J. J. RACH. 1999. Behavioral responses to disturbance in freshwater mussels with implications for conservation and management. Journal of the North American Benthological Society 18:381-390.
105. WALTER, M. T., M. F. WALTER, E. S. BROOKS, T. S. STEENHUIS, J. BOLL, AND K. WEILER. 2000. Hydrologically sensitive areas: variable source area hydrology implications for water quality risk assessment. Journal of Soil and Water Conservation 3:277-284.
106. WALTERS, C. J. 1986. Adaptive management of renewable resources. Macmillan Publishing, New York.
107. WATERS, T. F. 1995. Sediment in streams: sources, biological effects and control. American Fisheries Society Monograph 7:1-251.
108. WIEGAND, T., K. A. MOLONEY, J. NAVES, AND F. KNAUER. 1999. Finding the missing link between landscape structure and population dynamics: a spatially explicit perspective. American Naturalist 154:605-627.
109. WIENS, J. A. 1997. Metapopulation dynamics and landscape ecology. Pages 43-62 in I. A. Hanski and M. E. Gilpin (editors). Metapopulation biology: ecology, genetics, and evolution. Academic Press, San Diego, California.
110. WIENS, J. A. 2002a. Riverine landscapes: taking landscape ecology into the water. Freshwater Biology 47:501-515.
111. WIENS, J. A. 2002b. Central concepts and issues of landscape ecology. Pages 3-21 in K. J. Gutzwiller (editor). Applying landscape ecology in biological conservation. Springer, New York.
112. WILCOVE, D. S., D. ROTHSTEIN, J. DUBOW, A. PHILLIPS, AND E. LOSOS. 1998. Quantifying threats to imperiled species in the United States. BioScience 48:607-615.
113. WILKINSON, C. D., AND D. R. EDDS. 2001. Spatial pattern and environmental correlates of a Midwestern stream fish community: including spatial autocorrelation as a factor in community analyses. American Midland Naturalist 146:271-286.
114. WITH, K. A. 1994. Using fractal analysis to assess how species perceive landscape structure. Landscape Ecology 9:25-36.
115. WITH, K. A., AND T. O. CRIST. 1995. Critical thresholds in species' responses to landscape structure. Ecology 76:2446-2459.
116. WOOLNOUGH, D. A. 2006. The importance of host fish in long range transport of unionids in large rivers. PhD Thesis, Iowa State University, Ames, Iowa.
117. YOUNG, M. R., P. J. COSGROVE, AND L. C. HASTIE. 2001. The extent of, and causes for, the decline of a highly threatened naiad: Margaritifera margaritifera. Pages 337-357 in G. Bauer and K. Wächtler (editors). Ecology and evolution of the freshwater mussels unionoida. Springer-Verlag, Berlin, Germany.
118. ZIGLER, S., T. NEWTON, J. STEUER, M. BARTSCH, AND J. SAUER. 2008. Importance of physical and hydraulic characteristics to unionid mussels: a retrospective analysis in a large river reach. Hydrobiologia 598:343-360. DOI: 10.1007/s10750-007-9167-1.
119. ZWICK, P. 1992. Stream habitat fragmentation - a threat to biodiversity. Biological Conservation 1:80-87.