Drift and upstream colonisation
Acrive drift and passive dislodgement both result
in downstream movement of organisms (Secrion
2-3d), and one of rhe mosr enduring conundrums
in river ecology is why this does not resulr in
depletion of upstream popularions- Muller (1954) proposedthat this could be explained, for aquatic 1insects, by a colonisation cycle: Iarval movement is
essentially downsrream, so adults will fly
upstream to compensate for this. Unfornrnately,
the evidence available provides little supporr for
this hypothesis- Some studies (e.g. Elliom, 1967)
have found no evidence for a tendency ro fly
upstream and even those which do find supporr
for the idea, such as Flecker and Allan,s (1tSS)
study of mayflies in Colorado, are not as clear as
may be expected- These aurhors shorved a significant
tendency for Baetis spp. ro fly upstream,
although, in several of their samples, *o.. *...
flying downstream, whereas Rhitbrogena hageni
tended to fly dowrsrream- A further probtem wirh
the colonisarion cvcle hypothesis is that it does not
explain how species without a flying srage, such as
crustaceans, can persist in upstream stretchesUpstream
movemenrs by benthos moving over or
'r,vithin sedimenr (positive rheoraxis) have been
recorded - see Allan (1995) for a review - bur,
again, are inadequate to compensate for downstream
driftAllan
(1995) suggests thar upstream movement
is unnecessary to replace losses, as drift from
headrvaters is loss of surplus individuals and
enough u,ill remain ro replenish rhe popularionThe
evidence is, horverer, equivocal and, for the
moment, the phenomenon remains unexplained-