scholarly journals Selection Forces Driving Herding of Herbivorous Insect Larvae

2021 ◽  
Vol 9 ◽  
Author(s):  
Emma Despland

Herding behavior is widespread among herbivorous insect larvae across several orders. These larval societies represent one of several different forms of insect sociality that have historically received less attention than the well-known eusocial model but are showing us that social diversity in insects is broader than originally imagined. These alternative forms of sociality often focus attention on the ecology, rather than the genetics, of sociality. Indeed, mutually beneficial cooperation among individuals is increasingly recognized as important relative to relatedness in the evolution of sociality, and I will explore its role in larval insect herds. Larval herds vary in in the complexity of their social behavior but what they have in common includes exhibiting specialized social behaviors that are ineffective in isolated individuals but mutually beneficial in groups. They hence constitute cooperation with direct advantages that doesn’t require kinship between cooperators to be adaptive. Examples include: trail following, head-to-tail processions and other behaviors that keep groups together, huddling tightly to bask, synchronized biting and edge-feeding to overwhelm plant defenses, silk production for shelter building or covering plant trichomes and collective defensive behaviors like head-swaying. Various selective advantages to group living have been suggested and I propose that different benefits are at play in different taxa where herding has evolved independently. Proposed benefits include those relative to selection pressure from abiotic factors (e.g., thermoregulation), to bottom-up pressures from plants or to top-down pressures from natural enemies. The adaptive value of herding cooperation must be understood in the context of the organism’s niche and suite of traits. I propose several such suites in herbivorous larvae that occupy different niches. First, some herds aggregate to thermoregulate collectively, particularly in early spring feeders of the temperate zone. Second, other species aggregate to overwhelm host plant defenses, frequently observed in tropical species. Third, species that feed on toxic plants can aggregate to enhance the warning signal produced by aposematic coloration or stereotyped defensive behaviors. Finally, the combination of traits including gregariousness, conspicuous behavior and warning signals can be favored by a synergy between bottom-up and top-down selective forces. When larvae on toxic plants aggregate to overcome plant defenses, this grouping makes them conspicuous to predators and favors warning signals. I thus conclude that a single explanation is not sufficient for the broad range of herding behaviors that occurs in phylogenetically diverse insect larvae in different environments.

2020 ◽  
Author(s):  
Yadugiri V Tiruvaimozhi ◽  
Sumanta Bagchi ◽  
Mahesh Sankaran

AbstractArbuscular mycorrhizal fungal (AMF) symbioses with plants can be influenced by top-down forces such as grazing, and also by bottom-up forces such as soil resource availability, both of which are being altered by anthropogenic and global change drivers. While the influence of each of these factors on AMF symbioses has been widely studied, explicit tests of the relative strengths of top-down versus bottom-up influences on these ubiquitous plant root symbioses are few. We studied AMF colonization responses of four species of graminoids (3 grasses Elymus longae-aristatus, Leymus secalinus and Stipa orientalis, and a sedge Carex melanantha) common to semiarid high-altitude rangelands of the Spiti region, Trans-Himalaya, to changes in a top-down driver, grazing intensity (through short-term clipping and long-term grazer exclusion experiments), and a bottom-up driver, water availability (using irrigation treatments, and by evaluating responses to annual precipitation levels across years). Over three years, AMF colonization in all four host species was influenced by precipitation, with the highest and lowest AMF colonization levels corresponding to years with the lowest and highest rainfall, respectively. However, responses to long-term grazer exclusion differed among host species, and across years: while some species showed decreases in AMF colonization levels under grazing, others showed increases from ungrazed control levels, and these responses changed, even reversed, across years. Responses to short-term clipping and irrigation treatments also differed among hosts, with some species responding to irrigation alone, some to clipping and irrigation combined, and others showing no changes in AMF colonization from control levels in any of the treatments. In our study, long-term changes in water availability influenced AMF colonization levels, while short-term responses were host specific. Responses to above-ground tissue loss, however, differed among host species both in the long- and short-term. Overall, this study demonstrates that while AMF colonization levels correspond to annual precipitation levels in this semiarid ecosystem, host species also play a role in influencing plant-AMF interactions in these rangelands, with colonization levels and responses to abiotic factors changing with host species.


2021 ◽  
Vol 12 ◽  
Author(s):  
Rui Wu ◽  
Simcha Lev-Yadun ◽  
Lu Sun ◽  
Hang Sun ◽  
Bo Song

Glandular trichomes are well known to participate in plant chemical and physical defenses against herbivores, especially herbivorous insects. However, little is known about large-scale geographical patterns in glandular trichome occurrence. Herbivory pressure is thought to be higher at low elevations because of warmer and more stable climates. We therefore predicted a higher proportion of species with glandular trichomes at low elevations than at higher elevations. We compiled glandular trichome data (presence/absence) for 6,262 angiosperm species from the Hengduan Mountains (a global biodiversity hotspot in southwest China). We tested the elevational gradient (800–5,000 m a.s.l.) in the occurrence of plant species with glandular trichomes, and its correlations with biotic (occurrence of herbivorous insects) and abiotic factors, potentially shaping the elevational gradient in the occurrence of glandular trichomes. We found a significantly positive relationship between elevation and the occurrence of glandular trichomes, with the proportion of species having glandular trichomes increasing from 11.89% at 800 m a.s.l. to 17.92% at above 4,700 m. This cross-species relationship remained significant after accounting for phylogenetic relationships between species. Herbivorous insect richness peaked at mid-elevations and its association with the incidence of glandular trichomes was weak. Mean annual temperature was the most important factor associated negatively with glandular trichomes. Our results do not support the hypothesis that plant defenses decrease with increasing elevation. In contrast, a higher proportion of plant species with glandular trichome toward higher elevations is observed. Our results also highlight the importance of considering the simultaneous influences of biotic and abiotic factors in testing geographical variation in multifunctional plant defenses.


PeerJ ◽  
2020 ◽  
Vol 8 ◽  
pp. e8782
Author(s):  
Emma Despland ◽  
Paola G. Santacruz

The recent introduction in a tropical agricultural environment of a weedy open-habitat plant (Solanum myriacanthum) and subsequent host range expansion of a common forest-edge butterfly (Mechanitis menapis) onto that plant provides an opportunity to examine reconfiguration of tritrophic networks in human-impacted landscapes. The objectives of this study were (1) determine if the caterpillars on the exotic host are more or less limited by plant defenses (bottom-up forces) and if they experience enemy release (decrease of top-down pressure) and (2) define how anthropic open pasture habitat influences the herbivore’s tritrophic niche. Field and laboratory monitoring of larval survival and performance on a native (Solanum acerifolium) host plant and the exotic (S. myriacanthum) host plant were conducted in the Mindo Valley, Ecuador. Plant physical defenses were also measured. Results showed that larval mortality was mostly top-down on S. acerifolium, linked to parasitism, but mostly bottom-up on S. myriacanthum, possibly linked to observed increased plant defenses. Thus, in the absence of co-evolved relationships, herbivores on the exotic host experienced little top-down regulation, but stronger bottom-up pressures from plant defenses. These findings provide a rare empirical example of enemy-free space as a mechanism underlying host-range expansion. S. myriacanthum was less colonized in open pastures than in semi-shaded habitats (forest edges, thickets): fewer eggs were found, suggesting limited dispersal of adult butterflies into the harsh open environments, and the survival rate of first instar larvae was lower than on semi-shaded plants, likely linked to the stronger defenses of sun-grown leaves. These findings show how environmental conditions modulate the rewiring of trophic networks in heavily impacted landscapes, and limit a biocontrol by a native herbivore on an invasive plant in open habitats.


PsycCRITIQUES ◽  
2005 ◽  
Vol 50 (19) ◽  
Author(s):  
Michael Cole
Keyword(s):  
Top Down ◽  

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