In the southern Appalachian region of North America, the phylogenetically convergent shells of the polygyrid snails Triodopsinae Neohelix major (Binney) and Polygyrinae Mesodon normalis (Pilsbry) are even more convergent in size and shape in sympatry (7 sites) than in allopatry (23 and 10 sites). Environmental correlations account for 34% and 30% of size and shape variations in N. major (larger, taller, and more loosely coiled at northern, high-altitude, sheltered sites), but for only 14% and 9% in M. normalis (larger, flatter, and more loosely coiled at south-facing, exposed sites). The statistical significance of the sympatric convergence dropped out when these correlations were removed. This phenomenon helps account for the many cases in eastern North America of nearly identical land-snail shells in sympatry and questions the importance of competitive character displacement in the evolution of land-snail shell morphology. This apparently nonmimetic case of sympatric convergence provides an unusually precise and well-delimited, naturally replicated experiment in evolutionary morphology, which is analyzed for controlling factors in a follow-up paper.

In the southern Appalachian region of North America, the phylogenetically convergent shells of the polygyrid snails Triodopsinae Neohelix major (Binney) and Polygyrinae Mesodon normalis (Pilsbry) are even more convergent in size and shape in sympatry (7 sites) than in allopatry (23 and 10 sites). Environmental correlations account for 34% and 30% of size and shape variations in N. major (larger, taller, and more loosely coiled at northern, high-altitude, sheltered sites), but for only 14% and 9% in M. normalis (larger, flatter, and more loosely coiled at south-facing, exposed sites). The statistical significance of the sympatric convergence dropped out when these correlations were removed. This phenomenon helps account for the many cases in eastern North America of nearly identical land-snail shells in sympatry and questions the importance of competitive character displacement in the evolution of land-snail shell morphology. This apparently nonmimetic case of sympatric convergence provides an unusually precise and well-delimited, naturally replicated experiment in evolutionary morphology, which is analyzed for controlling factors in a follow-up paper.

A fresh assessment of reproductive behaviour and anatomy, combined with new allozyme data, results in a phylogenetic hypothesis for the 23 polygyrid genera that differs substantially from previous attempts. Analyses based on this phylogeny suggest that (a) evolutionary transition from internal to external sperm exchange occurred only once in polygyrids, with unusual functional intermediates still extant; (b) polygyrid biogeographic history paralleled that of plethodontid salamanders, with hypothesized vicariance events at about 145, 120, 65 and 40 MA, and, dispersal events at about 55 and 40 MA; and (c) iterative shell evolution occurred despite a phylogenetic constraint on ontogenetic whorl-expansion rate.

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Madagascar's magnificent and environmentally threatened endemic radiation of the land-snail genus Tropidophora Troschel has recently been classified into three subgenera, 95 species and 142 varieties, based on often subtle conchological variation among small samples; it seems best to ignore temporarily this confusing plethora of names until true biological species and their relationships are better understood. The author's field work in 1990 succeeded in obtaining live Tropidophora from 40 populations distributed throughout the island. Allozyme analysis (108 snails, 15 loci, 117 alleles) yielded a cladogram suggesting nine genetically coherent taxa (T. taxa A-I), each represented in the collections by 1-10 populations.Comparisons among shells (total 1634) and penes (total 31 from 20 populations representing eight of the nine taxa) revealed: (1) two conchologically indistinguishable taxa (H and I) fixed for alternative alleles at 13 of 1Fi loci, with only a very subtle difference in penes, and with mosaic and overlapping geographical distributions in the Northeast; (2) two extremely polytypic taxa (C in the Southwest, F in the Southeast) with parallel trends toward depressed, broadly umbilicate, heavily sculptured shells with apertural lips widely reflected at the umbilicus at inland, more arid sites, resulting in sympatric convergence; (3) one southeastern taxon (G) in which the penis apparently doubles in length but the shell does not change 6.0 km to the northwest, but in which the shell shifts dramatically in sculpture, colour, and lip reflection 0.5 km to the westnorthwest; and (4) generally such extreme intra-and interpopulation variation in shell and male-genital characters as to render many of them dubious at best for systematics. Thus the Madagascan Tropidophora present a fascinatingly complex problem in evolutionary morphology/ecology, the solution of which will require even more extensive collecting, followed by molecular comparisons or detailed anatomical comparisons, or both. The total number of biological species is still likely to be quite large, despite irrelevance of much of the current taxonomy, because many smaller species remain to be discovered. 0 1995 The Linnean Society of London ADDITIONAL