. Not surprisingly, speculation is rife about its evolutionary genesis. Some favor the hygric hypothesis (reduction of respiratory water loss; see review by Lighton, 1996) and others the chthonic hypothesis (decoupling of O 2 and CO 2 exchange, and enhanced ability to exchange gases in hypercapnic and hypoxic environments; Lighton, 1996). Recently Hetz and Bradley (2005) proposed an ingenious third theory, namely avoidance of O 2 toxicity: the insect guards its interior against the toxic effects of oxygen by restricting access to its tracheal system, particularly in the C (closed-spiracle) phase and later in the F (fluttering-spiracle) phase of the DGC, during which internal P O 2 is regulated at <4·kPa (see reviews by Kestler, 1985;Lighton, 1996Lighton, , 1998Chown and Nicolson, 2004;. We refer below to any spiracle-control strategies that limit internal P O 2 as oxygen guarding.The Hetz-Bradley hypothesis is tenable if, in fact, the DGC creates a milieu interieur with a lower mean P O 2 than alternative means of gas exchange. But paradoxically, a delayed oxygen exposure penalty is created while P O 2 is maintained at a low level of ca. 4·kPa during the DGC's F phase. During the F phase, thanks to stringent spiracular control, CO 2 escapes at <25% of production rate (Lighton, 1988 and references therein). Periodic flushing of this accumulated CO 2 is therefore necessary, and in fact this flushing -known as the open-spiracle (O) or burst phase -is in large part the operational definition of the DGC. As a sideeffect of flushing the CO 2 produced during the C and F phases, in the O phase internal P O 2 rises to near-atmospheric levels throughout the tracheal system (Hetz and Bradley, 2005 and references therein).Obtaining comparative data that bear on the oxygenguarding hypothesis is difficult. While it is interesting that Drosophila endowed with extra superoxide dismutase and catalase copies show a small but significant increase in lifespan (see review by Sohal et al., 2002), (a) the effect is minimal if wild-type, rather than genetically compromised, flies are used in the experiments (ibid), (b) overexpression of antioxidants may actually have deleterious effects (ibid), and (c) most oxygen toxicity experiments are short-term and make the assumption that evolutionary fitness and individual longevity are coterminous. Thus current oxygen toxicity data are not The ability of some insects to engage in complex orchestrations of tracheal gas exchange has been well demonstrated, but its evolutionary origin remains obscure. According to a recently proposed hypothesis, insects may employ spiracular control of gas exchange to guard tissues against long-term oxidative damage by using the discontinuous gas-exchange cycle (DGC) to limit internal oxygen partial pressure (P O 2). This manuscript describes a different approach to oxygen guarding in the lower termite Zootermopsis nevadensis. These insects do not display a DGC but respond to elevated oxygen concentrations by restricting spiracular area, resulting in a transient ...