Peter O'Byrne wrote:

>Things do not look good for the "one orchid species, one insect species"
>hypothesis.

Indeed. But two points that you may have elided:

1: The complexity / zygomorphy / whatever of the orchid flower is not un
caprice de dieu, but something that is actively evolving under insect control.
Ergo, how it looks at microscopic detail matters to insects. I suspect that
the flower at least sometimes and to a degree "imprints" the pollinator - in
effect, says 'was it good for you too? Well, go and find another just like
me'. THis, because the pollinia system means that all of the gametes are bound
up in one (all right, up to eight) package(s), but in effect the flower gets
just one shot. Other insect pollinators tend to have dusty pollen, many
anthers, lots of bites at the bug's cherry. Orchids are less of a shotgun,
more a precise sniper rifle and they get just one shot. It has to hit its
target first time, which means that the insect in question has to be sent on a
quest through the forest. My point: floral form may be a self-driving
evolutionary isolation barrier, operating through insect 'psychology'.

2: Two clades diverge by many small steps, and what looks like a huge
phenotypic change may be a point mutation in a e.g. homeobox gene. However,
what makes for evolutionary isolation or inclusion is always the phenotype.
Thus one could have a swarm - Arundina - building clade-potential but still
apparently interbreeding, despite being anything from a harebell to a
hollyhock in appearance, whilst others may be separated by a single whisker or
callus. Coelogyne, for example - the Clayton book shows many species which are
all-but identical but which apparently operate in sexual isolation, perhaps
due to allopatry - with all of the problems that you mention - with changed
time of flowering (in my view, an undervalued variant in seasonal regions) or
perhaps due to sensitivities amongst our friends, the bugs; as (1).

One should note that "isolation" is anyway a statistical rather than a binary
concept. If the chances of reproduction between interbreeding set A and set B
are 80% of the rate of breeding within the set, then the progeny will always
be fewer than those of either A or B, and thus at a major disadvantage. This
could be offset by hybrid vigor, perhaps, but not always; particularly if A
and B are closely related and have much the same recessive burden. .
_____________________________________
Oliver Sparrow
Tel: UK (0)20 7736 9716
www.chforum.org
www.treknepal.org
www.datafreeze.com
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