336 HETEROCHRONY AND PAEDOMORPHOSIS
drozoans), Mystacocarida (Crustacea), and Acochlidiacea (Mollusca).
It is a testimony to the importance of heterochrony that all these
groups have received a progenetic interpretation. The Actinulida are
generally regarded as progenetic (Hadzi, 1963, p. 94). Hessler (1971,
p. 87) has urged a progenetic interpretation of the Mystacocarida in
opposition to previous claims for their "primitive" nature. Riedl
(1969, p. 856) points to the paedomorphic features of gnathostomulids in suggesting their possible progenetic origin from much
larger fossil conodonts. Odhner (1952) suggests that the family Microhedylidae of the Acochlidiacea may be progenetic.
The Role of Heterochrony in Macroevolution:
Contrasting Flexibilities for Progenesis and Neoteny
Progenesis
Progenesis has usually been dismissed as an agent of degeneration
with no evolutionary importance. I have tried to rescue it from this
charge by demonstrating its immediate significance for precocious
maturation in r-selected regimes (and, in some cases, for small body
size). But de Beer's dismissal was more an assessment of retrospective
significance in macroevolution than a denial of adaptive value in local
environments.
The link of progenesis to r selection might seem to affirm the usual
denial of retrospective significance. "Evolutionary importance" is an
ambiguous notion (to put it mildly); in conventional writing about
macroevolution, a process partakes of this recondite "importance"
when it contributes to the classical material of macroevolutionary success: slow, continuous, and sustained trends toward more complex
morphology and greater diversity. These trends are the material of K
selection; r selection, with its emphasis on production of offspring at
the expense of perfection in morphology, can only serve as a brake
upon such "progress." As Dobzhansky wrote in his prophetic article of
1950, long before theoretical ecology codified the concept: "Physical
factors, such as excessive cold or drought, often destroy great masses
of living beings, the destruction being largely fortuitous with respect
to the individual traits of the victims and the survivors . . . Indiscriminate destruction is countered chiefly by development of increased fertility and acceleration of development and reproduction,
and does not lead to important evolutionary advances" (p. 220; see
also Murphy, 1968, p. 402). Levin has noted the restriction of recombination in r-selected plants, K-selected genomes, he argues, do not
congeal because pathogens and herbivores exert intense pressure and