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associated with the latter (Clausen 1954 ; Nygren 1966 ; DeWet and Stalker 1974 ;
Asker 1979 ; Catling 1981, 1982 ). When compared with autogamy, agamospermy
requires less extreme morphological adaptation (Ornduff 1969 ) and produces no
inbreeding depression (Charlesworth and Charlesworth 1979 ). At the same time,
fertility is assured; the potential for rapid colonization is retained; heterozygosity is
maintained; and the fixation of particular patterns of variability (new adaptive gene
combinations) may be facilitated (Baker 1955 ; Antonovics 1968 ; Jain 1976 ; Solbrig
1976 ; Marshall and Weir 1979 ; Catling 1980b, 1982 ; Lloyd 1988 ). Short-term fit-
ness may, thus, be combined with the advantages of genetic recombination (Catling
1982 ). The latter includes, for example, the ability to recombine new, advantageous
mutants, permitting evolutionary responses to changing environmental conditions;
the ability of the best-adapted genotypes to escape the accumulation of linked, dis-
advantageous but nonlethal mutants (“Muller's Ratchet”); and the ability to fill the
maximum number of environmental niches through a high level of genetic variation
among individuals.
Agamospermy is also thought to confer advantages over sexual reproduction in
certain areas, where it may reflect adaptation to isolation or to a reduction in the
availability or activity of pollinators (Lloyd 1980, 1988 ; Manning 1981 ; Catling
1982 ; Schmidt and Antlfinger 1992 ). Such sites include wide expanses of recently
glaciated territory, edges of a species range, recently established vegetation zones,
or more or less isolated bioclimatic zones (e.g., Bayer and Stebbins 1980 ; Catling
1982 ; Schmidt and Antlfinger 1992 ). For example, the three species, S. magnicamp-
orum, S. ochroleuca , and S. odorata , which appear to be sexual over most of their
distributions, reveal an apparent association between agamospermy and range lim-
its, particularly northeastern range limits (Catling 1982 ). S. casei is distributed
within formerly glaciated territory (Prest 1970 ; Ives et al. 1975 ; Catling 1990 ), and
S. casei var. novascotiae lies in the relatively isolated bioclimatic zone of southern
Nova Scotia (Fernald 1921 ; Roland and Smith 1962 -1969; Catling 1981, 1982,
1990 ). The association of agamospermy with northeastern range limits, glaciated
territory, and isolated bioclimatic zones, its augmentation of other isolating mecha-
nisms, and its presence in a complex of closely related taxa suggest that it has been
a significant factor in the production of new species (Catling 1982, 1983a ).
Sexual Species or Populations
This section examines sexual reproduction in uniformly sexual species of Spiranthes
and the facultatively agamospermous populations of the S. cernua complex whose
asexual reproduction was discussed above. Uniformly sexual species include: S. lacera
(Rafinesque) Rafinesque var. lacera (northern slender ladies'-tresses), S. lacera var.
gracilis (Bigelow) Luer (southern slender ladies'-tresses), S. tuberosa Rafinesque (lit-
tle ladies'-tresses), S. vernalis Engelmann and Gray (spring ladies'-tresses), S. lacini-
ata (Small) Ames (lace-lip ladies'-tresses), S. romanzoffiana Chamisso (hooded
ladies'-tresses), S. diluvialis Sheviak (Ute ladies'-tresses), S. lucida (H. H. Eaton)
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