UNIVERSIDAD DE LA AMAZONIA

FACULTAD DE INGENIERÍA
PROGRAMA DE INGENIERÍA AGROECOLÓGICA
CURSO DE ENTOMOLOGÍA

LECTURA 3

Fuente:
Gullan, P. J.; Cranston, P. S. 2014. The Insects. An outline of Entomology. Editorial
Chapman & Hall. Quinta Edición. Estados Unidos de América. 624 p.

THERE WERE GIANTS – EVOLUTION OF INSECT GIGANTISM

We are used to insects being amongst the smaller animals we encounter, with many
being inconspicuous in terms of size and lifestyles. However, back in the
Carboniferous and Permian, some were very large, exemplified by the giant Bolsover
dragonfly and a palaeodictyopteran on a Psaronius tree fern, depicted here (inspired
by a drawing by Mary Parrish in Labandeira 1998). Late Palaeozoic Ephemeroptera
and dragonfly-like Meganisoptera had wingspans of up to 45 cm and 71 cm,
respectively. Gigantism at this time was evident, not only among insects but also in
branchiopod crustaceans, certain other invertebrates, and some amphibians.

From the earliest recognition of gigantism in the insect fossil record, elevated
atmospheric oxygen levels (above today’s 20%) were suggested, although the
geological processes responsible were unknown. Restriction of modern insects to a
modest maximum size and shape (volume) seems to derive from the physical laws
governing gas transport along the tracheae to the sites of uptake and release in cells.
Flight is so highly demanding of oxygen that a large, winged insect in a modern
oxygen atmosphere would require such dense tracheae that there would be inadequate
space for the muscles and other internal organs.
 UNIVERSIDAD DE LA AMAZONIA
FACULTAD DE INGENIERÍA
PROGRAMA DE INGENIERÍA AGROECOLÓGICA
CURSO DE ENTOMOLOGÍA

New biogeochemical models infer elevated atmospheric oxygen during the

Carboniferous and Permian as shown in the graph. Oxygen levels of at least 30%,
half as much again above today’s levels, imply a very different (and more flammable)
world in the late Carboniferous to the mid-Permian. For insects, higher oxygen
pressures promoted greater oxygen diffusion via tracheae, and reduced the demand
for the thorax to be packed with tracheae to power flight. Furthermore, with elevated
oxygen, air would be denser, further facilitating flight. This hypothesis for Palaeozoic
gigantism has obvious appeal, although morphological and physiological
consequences (including ability to actively ventilate tracheae) of alterations of
gaseous composition need further study.

The drop in atmospheric oxygen from the mid-Permian onwards coincided with
reduction in maximum size of winged insects, as expected. This trend continued
through to the end-Permian mass extinction, when many insects became extinct,
including those groups in which gigantism had been prevalent. However, the decrease
in maximum size from the Permian to modern times seems too large to be explained
by declining atmospheric oxygen alone. Furthermore, the postulated recurrence of
high oxygen in the Cretaceous and early Tertiary was not associated with regain of
large-sized insects. The explanation favoured for the subsequent disconnect between
atmospheric oxygen levels and insect body size was the arrival of flying vertebrate
predators, namely birds and bats.

These insectivores would have competed with larger insects, over-riding the effect of
environmental change on body size. The air was no longer the exclusive preserve of
winged insects. Thus, since the Tertiary, maximum sizes attained for insects such as
dragonflies and mayflies are below the upper limits of their extinct distant ancestors,
and less than their gas-exchange physiology would permit.

Pregunta de reflexión:

1. ¿Por qué los primeros insectos que poblaron el planeta Tierra eran de grandes
tamaños y, los que existen actualmente son de pequeños tamaños?