The origin of tetrapod herbivory: effects on local plant diversity

The origin of herbivory in the Carboniferous was a landmark event in the evolution of terrestrial ecosystems, increasing ecological diversity in animals but also giving them greater influence on the evolution of land plants. We evaluate the effect of early vertebrate herbivory on plant evolution by comparing local species richness of plant palaeofloras with that of vertebrate herbivores and herbivore body size. Vertebrate herbivores became diverse and achieved a much greater range of body sizes across the Carboniferous–Permian transition interval. This coincides with an abrupt reduction in local plant richness that persists throughout the Permian. Time-series regression analysis supports a negative relationship of plant richness with herbivore richness but a positive relationship of plant richness with minimum herbivore body size. This is consistent with studies of present-day ecosystems in which increased diversity of smaller, more selective herbivores places greater predation pressures on plants, while a prevalence of larger bodied, less selective herbivores reduces the dominance of a few highly tolerant plant species, thereby promoting greater local richness. The diversification of herbivores across the Carboniferous–Permian boundary, along with the appearance of smaller, more selective herbivores like bolosaurid parareptiles, constrained plant diversity throughout the Permian. These findings demonstrate that the establishment of widespread vertebrate herbivory has structured plant communities since the late Palaeozoic, as expected from examination of modern ecosystems, and illustrates the potential for fossil datasets in testing palaeoecological hypotheses.

The nature of open ecosystems

If open ecosystems were of recent anthropogenic origin, linked to human activity in the last millennia, they should support an impoverished biota assembled in large part from forest-dwelling species. Yet several of the world’s biodiversity hotspots are open ecosystems, rich in species and rich in endemics. This chapter introduces the diversity of open ecosystems and the distinction between old growth and secondary, early-successional vegetation. The functional traits of species in open ecosystems can be highly informative as to the dominant consumers maintaining their structure. Traits adapted to different fire regimes and those adapted to vertebrate herbivory are considered and compared. Faunal differences between open and closed vegetation are beginning to emerge from both invertebrate and vertebrate studies and indicate the importance of vegetation structure for habitat choice.

Open Ecosystems

This book explores the geography, ecology, and antiquity of ‘open ecosystems’, which include grasslands, savannas, and shrublands. They occur in climates that can support closed forest ecosystems and often form mosaics with forest patches. With the aid of remote sensing, it is now clear that open ecosystems are a global phenomenon and occur over vast areas in climates that could also support forests. This book goes beyond regional narratives and seeks general explanations for their existence. It develops the theme of open ecosystems as being widespread and ancient, with a distinct biota from that of closed forests. It examines hypotheses for their maintenance in climate zones favouring the development of forests, including soils hostile for tree growth, fire, and vertebrate herbivory.

Vertebrate florivory of vascular epiphytes: the case of a bromeliad

The avoidance of vertebrate herbivory is thought to be one of the possible drivers for the evolution of epiphytism. Scarce literature suggests that epiphyte herbivory is mainly related to insect attack on reproductive structures. In a pine-oak forest we observed almost all inflorescences of an epiphytic bromeliad (Tillandsia carlos-hankii) with signs of florivory; the degree of damage suggested that vertebrate herbivores could be involved. To assess the intensity of vertebrate florivory damage we recorded the percentage of damaged individuals in a 500 m2 plots during two flowering seasons. To identify possible vertebrate herbivores, we installed 20 mixed capture stations, 10 photo-traps focused on bromeliads and analyzed stomach contents of captured vertebrates. Florivory was observed on 62% of individuals during the first flowering season and 77% on the second; and average one individual lost 41% of reproductive structures. Vertebrates associated with florivory were a bird, Icterus bullockii (Aves, Passeriformes, Icteridae), a squirrel Sciurus aureogaster (Mammalia, Rodentia, Sciuridae), and mice, Peromyscus gratus, P. levipes and P. aztecus (Mammalia, Rodentia, Cricetidae). Our results suggest that vascular epiphytes are used as opportunistic resources for small vertebrates during seasons when preferred resources are scarce.

Benefits of jasmonate-dependent defenses against vertebrate herbivores in nature

Endogenous jasmonates are important regulators of plant defenses. If and how they enable plants to maintain their reproductive output when facing community-level herbivory under natural conditions, however, remains unknown. We demonstrate that jasmonate-deficient Nicotiana attenuata plants suffer more damage by arthropod and vertebrate herbivores than jasmonate-producing plants in nature. However, only damage by vertebrate herbivores translates into a significant reduction in flower production. Vertebrate stem peeling has the strongest negative impact on plant flower production. Stems are defended by jasmonate-dependent nicotine, and the native cottontail rabbit Sylvilagus nuttallii avoids jasmonate-producing N. attenuata shoots because of their high levels of nicotine. Thus, endogenous jasmonates enable plants to resist different types of herbivores in nature, and jasmonate-dependent defenses are important for plants to maintain their reproductive potential when facing vertebrate herbivory. Ecological and evolutionary models on plant defense signaling should aim at integrating arthropod and vertebrate herbivory at the community level.