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took longer to flower than clonal ramets. Nevertheless, a variable but usually high
number of capsules were usually produced (Correll 1978 ; Brackley 1985 ; Homoya
1993 ; Reddoch and Reddoch 1997 ). The percentage of flowering rhizomes in each
patch usually ranged from 0 to 8%. However, at 2-4 year intervals the percentage
increased to over 8% and up to 30%. Although rhizomes were no longer connected
to one another, the intermittent large flowerings were synchronized within patches,
between patches, and between populations. Reddoch and Reddoch ( 2007 ) corre-
LATEDTHISSYNCHRONYWITHTHEOCCURRENCEOFANEXTENDEDWARMDRYPERIODIN-AYOF
the preceding year.
Kallunki ( 1981 FOUNDWIDEINTRASPECIlCVARIATIONINFRUITSETINFOURMIXED
open-pollinated populations of G. tesselata and G. oblongifolia from Michigan
(Table 1.2 ). A fruit set of 90% reported by Kipping ( 1971 ) for a small population of
G. oblongifolia IN#ALIFORNIAEXTENDSTHERANGEEVENFURTHER+ALLUNKIATTRIBUTEDTHE
variation she observed to differences in the abundance and visibility of Goodyera
SPECIESTHEPRESENCEORABSENCEOFCOMPETINGSPECIESANDORDIFFERENTIALEXPOSURE
to strong winds and human activity. In any case, mean fruit set values in naturally
pollinated plants of G. tesselata , G. repens , and G. oblongifolia were markedly
lower than in hand-pollinated plants (Table 1.2 ), suggesting that pollinator recruit-
ment might be a limiting factor. Flowering plants also produced a higher mean
number of new rhizomatous growths than nonflowering plants (Ackerman 1975 ),
implying an absence of resource limitation. Long-term studies are needed, however,
to sort out possible initial differences in plant vigor.
In Ackerman's ( 1975 ) study of G. oblongifolia in California, pollinaria were
REMOVEDFROMOFOPENPOLLINATEDmOWERSANDDEPOSITEDONOFEXPOSED
stigmas. Mature capsules developed in 46% of the flowers. Ackerman considered
the mortality rate, taken as the difference between the number of flowers pollinated
and the number of capsules that matured, to be significant. Intervals between polli-
nation and fertilization and between fertilization and maturation of the capsules
were short. Seeds were sometimes liberated within a few weeks of flowering (Catling
1990 ). Although Swamy ( 1949 ) thought such short intervals represented a primitive
condition, Ackerman ( 1975 ) believed they were related to mortality rates, repre-
senting a secondary adaptation to environmental conditions.
Wind distribution of the seeds and adaptation to common and widely occurring
pollinators indicate a potential for wide dispersal (Muesebeck et al. 1951 ; Ackerman
1975 ). Ackerman ( 1975 ) suggests that the disjunct population structure of G.
oblongifolia may reflect this potential. However, Case ( 1987 ) believes that, at least
in the Great Lakes region, disjunct populations may represent remnants of a once
more widespread distribution fragmented by glaciation.
A number of factors, therefore, appear to contribute to the reproductive success of
one or more species of Goodyera (Ackerman 1975 ). Seed production is increased by
self-compatibility, clonal growth, aggregation of flowers, nectar production, and adap-
tation to common pollinators; maintenance of genetic variability results from occa-
SIONALOUTCROSSINGPROMOTEDBYPROTANDRYANDHYBRIDIZATIONISRESTRICTEDTOANEXTENT
that permits individual species to maintain their identities; capsule mortality may
be minimized by short maturation time; and the establishment of new populations is
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