The relationship between toughness and herbivory is complex; despite the negative findings of some recent authors for dicots we hypothesize that either greater toughness or late folding can protect monocot leaves against herbivorous insects in tropical lowland rain forest, and that the relative importance varies widely with species. The difficulties of establishing unequivocally the roles of leaf toughness and leaf folding or rolling in a given case are discussed.

It is predicted that monocots will be found to experience lower rates of herbivory by invertebrates than dicots. The tough monocot leaves include both stiff leaves containing relatively little water at saturation (e.g. palms), and leaves which lack stiffness, are rich in water at saturation and roll readily during dry weather or even in bright sun around midday (e.g. gingers, heliconias and marants). Monocot leaves also show that it is possible for leaves to be notably tough throughout the expansion phase of development, something never recorded for dicots. The need to broaden the botanist's mental picture of a 'tough leaf' is emphasized.

In tropical lowland rain forest, we find that species with the leaves tightly folded or rolled until they reach at least 50% of final length occur in 10 of the 15 monocot families with >100 species, and in 12 of the 24 monocot families with <100 species, but in only seven of the 212 dicot families (eudicots and magnoliids). Earlier researchers have described how examples of tightly folded and rolled leaves develop, but most have not considered the potentially adaptive value of this pattern of growth. We hypothesize that it is a protection against herbivorous invertebrates. For tropical and temperate dicots, the young leaves have been found to suffer much smaller losses to herbivores while folded and rolled than after they are unfolded or unrolled. Being folded or rolled until a late stage involves an 'opportunity cost' in the loss of photosynthesis. Among dicots, defences involving such a cost (notably late development of photosynthetic systems in pendent soft young leaves) are typical of shade-tolerant species, which have longer-lived leaves than light-demanders. In contrast, among monocots late folding and rolling are found in both shade-tolerators and light-demanders. We hypothesize that late folding and rolling bring a net advantage to monocots in general, whether shadetolerant or light-demanding, despite the opportunity cost, because they mostly have fewer leaves per plant of a given size, and therefore an individual leaf is relatively more valuable to the plant. As a coda, we suggest that the 'sleep movements' of some tropical plants, and the circinate vernation of ferns and some cycads, provide protection against invertebrate herbivores through the apposition of two or more layers of leaf.

No abstract

This study examined the effects of insect herbivory on growth and mortality of seedlings of a mid‐ successional rainforest tree, Alphitonia whitei Braid. Two experiments were conducted in which seedlings were exposed to 0% and 50% natural defoliation by insect herbivores and placed in light gaps in simple notophyll vine forest at Paluma, near Townsville, North Queensland. In the first experiment, insect herbivory significantly increased mortality of 2‐month‐old seedlings. Smaller seedlings had significantly greater mortality rates than larger seedlings, irrespective of herbivory. A significantly greater proportion of smaller seedlings died from being smothered by fallen leaves and soil as a result of digging by vertebrates than for larger seedlings. In the second experiment, the effects of seedling age were examined by comparing 2‐month and 4‐month‐old seedlings. Mortality rates were significantly influenced by seedling age, with eight times greater survival of older seedlings than of younger seedlings. Although insect herbivory was correlated with a significant decrease in shoot mass and a significant increase in root:shoot ratio, there was no effect of insect herbivory on seedling survival in the second experiment. Thus, mortality rates were greater for seedlings if they were young or small (which indirectly results from insect herbivory), because small, young seedlings were more susceptibile to other mortality factors, such as burial by fallen debris and digging by vertebrates.