Biology Reference
In-Depth Information
as already noted the flowers of S. magnicamporum have a strong and distinctive
fragrance (Sheviak 1973, 1976, 1982 ). Catling and Brown ( 1983 ) think that pollina-
tors in mixed populations of these species may very well respond to these differ-
ences. At the same time, a general similarity in floral morphology may serve to
attract and reinforce the behavior of pollen vectors as in Goodyera (Kallunki 1981 ;
Catling and Brown 1983 ).
Pollinators and Pollination Mechanisms
The primary pollinators of this genus are medium sized (ca. 9-mm long) to com-
paratively large (15-20-mm long), long-tongued bees, particularly species of
Bombus and to lesser extent members of the Megachilidae (Tables 2.3-2.9 ) (Darwin
1862 ; Robertson 1893, 1929 ; Ames 1921 ; Godfery 1931, 1933 ; Catling 1980b,
1983c ; Sheviak 1982 ). Smaller halictine bees (species of Lasioglossum ( Dialictus
Robertson) and Augochlorella ) are adapted to the pollination of S. lucida (Table 2.7 );
visits of these bees to other species are often erratic and associated with inefficient
pollen transfer (see below) (Catling 1980b, 1983c ). Among the few remaining tabu-
lated insects, the importance of the andrenid Calliopsis andreniformis as a pollina-
tor of S. lacera (Table 2.5 ) remains to be determined. The same may be said of the
long-tongued bees of the genus Anthophora and the typhiid wasp, Myzinum , reported
as pollinators of S. diluvialis (Table 2.8 ). Sipes and Tepedino ( 1995 ) only rarely
observed them bearing any pollinaria, and they have not been implicated in the pol-
lination of other species of Spiranthes (Catling 1983c ).
In most Spiranthes , the nectar glands (calli) secrete their nectar into the base of
the floral tube (Fig. 2.5a ) (Ames 1921 ; Correll 1978 ; Catling 1982, 1983c ). The
relatively long, curved galea and projecting tongue of Bombus species and
Megachilidae are well-adapted to reach this nectar source. The galea is hinged to the
Stipes and when extended it reaches well forward of the head. As the insect inserts
its head into the flower to obtain the nectar, it brings the dorsal surface of the flat-
topped galea into contact with the viscidium (Fig. 2.5b ) (Catling 1983c ). The vis-
cidium is elongated and rigid in all northeastern species of Spiranthes , except S.
lucida (Fig. 2.3 ) (see below), and attaches readily to the stiff, flat-topped galea
(Fig. 2.5c ) (Catling 1980b, 1983c ; Catling and Catling 1991 ). It is oriented parallel
to the long axis of the proboscis and adheres to the middle or proximal upper surface
in bees with a short galea and to the distal upper surface in bees with a long galea.
A similar mode of attachment was reported for Bombus pollinators of S. romanzof-
fiana and S. diluvialis (see below) in western North America (Tables 2.6 and 2.8 )
(Larson and Larson 1987 ; Sipes and Tepedino 1995 ) and S. spiralis in England
(Darwin 1862 ; Godfery 1933 ).
The flowers open first near the base of the spike, and blooming proceeds upward
in sequence (e.g., Darwin 1862 ; Catling 1983c ; Sipes and Tepedino 1995 ). Individual
flowers are protandrous. In newly opened flowers, the lip and column are close
together and the stigma hidden (Fig. 2.4a ). At this male stage, incoming pollinia
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