Vincent Bonhomme

Slippery or sticky? Functional diversity in the trapping strategy of Nepenthes carnivorous plants

The pitcher-shaped leaves of Nepenthes carnivorous plants have been considered as pitfall traps that essentially rely on slippery surfaces to capture insects. But a recent study of Nepenthes rafflesiana has shown that the viscoelasticity of the digestive fluid inside the pitchers plays a key role. Here, we investigated whether Nepenthes species exhibit diverse trapping strategies. We measured the amount of slippery wax on the pitcher walls of 23 taxa and the viscoelasticity of their digestive liquid and compared their retention efficiency on ants and flies. The amount of wax was shown to vary greatly between species. Most mountain species exhibited viscoelastic digestive fluids while water-like fluids were predominant in lowland species. Both characteristics contributed to insect trapping but wax was more efficient at trapping ants while viscoelasticity was key in trapping insects and was even more efficient than wax on flies. Trap waxiness and fluid viscoelasticity were inversely related, suggesting the possibility of an investment trade-off for the plants. Therefore Nepenthes pitcher plants do not solely employ slippery devices to trap insects but often employ a viscoelastic strategy. The entomofauna specific to the plants habitat may exert selective pressures, favouring one trapping strategy at the expense of the other.

The plant-ant Camponotus schmitzi helps its carnivorous host-plant Nepenthes bicalcarata to catch its prey

The Bornean climber, Nepenthes bicalcarata , is unique among plants because it is both carnivorous and myrmecophytic, bearing pitcher-shaped leaves and the ant Camponotus schmitzi within tendrils. We explored, in the peat swamp forests of Brunei, the hypothesis that these ants contribute to plant nutrition by catching and digesting its prey. We first tested whether ants increased plants capture rate. We found that unlike most plant-ants, C. schmitzi do not exhibit dissuasive leaf-patrolling behaviour (zero patrol on 67 pitchers of 10 plants) but lie concealed under pitcher rim (13 ± 6 ants per pitcher) allowing numerous insect visits. However, 47 out of 50 individuals of the largest visitor dropped into the pitchers of five plants were attacked by ants and the capture rate of the same pitchers deprived of their ambush hunting ants decreased three-fold. We then tested whether ants participated in plants digestion. We showed in a 15-d long experiment that ants fed on prey and returned it in pieces in seven out of eight pitchers. The 40 prey deposited in ant-deprived pitchers remained intact indicating a weak digestive power of the fluid confirmed to be only weakly acidic (pH ~5, n = 67). The analysis of 10 pitcher contents revealed that prey, mainly ants and termites, was very numerous (~400 per pitcher per plant) and highly fragmented. Altogether, these data suggest a positive effect of C. schmitzi on both prey intake and breakdown. This ant–plant interaction could thus be a nutritional mutualism involving the unusual association of carnivory and myrmecotrophy.

Different pitcher shapes and trapping syndromes explain resource partitioning in Nepenthes species

Abstract Nepenthes pitcher plants display interspecific diversity in pitcher form and diets. This species‐rich genus might be a conspicuous candidate for an adaptive radiation. However, the pitcher traits of different species have never been quantified in a comparative study, nor have their possible adaptations to the resources they exploit been tested. In this study, we compare the pitcher features and prey composition of the seven Nepenthes taxa that grow in the heath forest of Brunei (Borneo) and investigate whether these species display different trapping syndromes that target different prey. The Nepenthes species are shown to display species‐specific combinations of pitcher shapes, volumes, rewards, attraction and capture traits, and different degrees of ontogenetic pitcher dimorphism. The prey spectra also differ among plant species and between ontogenetic morphotypes in their combinations of ants, flying insects, termites, and noninsect guilds. According to a discriminant analysis, the Nepenthes species collected at the same site differ significantly in prey abundance and composition at the level of order, showing niche segregation but with varying degrees of niche overlap according to pairwise species comparisons. Weakly carnivorous species are first characterized by an absence of attractive traits. Generalist carnivorous species have a sweet odor, a wide pitcher aperture, and an acidic pitcher fluid. Guild specializations are explained by different combinations of morpho‐functional traits. Ant captures increase with extrafloral nectar, fluid acidity, and slippery waxy walls. Termite captures increase with narrowness of pitchers, presence of a rim of edible trichomes, and symbiotic association with ants. The abundance of flying insects is primarily correlated with pitcher conicity, pitcher aperture diameter, and odor presence. Such species‐specific syndromes favoring resource partitioning may result from local character displacement by competition and/or previous adaptations to geographically distinct environments.

Visual field shape and foraging ecology in diurnal raptors

ABSTRACT Birds, particularly raptors, are believed to forage primarily using visual cues. However, raptor foraging tactics are highly diverse – from chasing mobile prey to scavenging – which may reflect adaptations of their visual systems. To investigate this, we studied the visual field configuration of 15 species of diurnal Accipitriformes that differ in such tactics, first focusing on the binocular field and blind area by using a single-traits approach, and then exploring the shape of the binocular field with a morphometric approach. While the maximum binocular field width did not differ between species with different foraging tactics, the overall shape of their binocular fields did. In particular, raptors chasing terrestrial prey (ground predators) had a more protruding binocular field and a wider blind area above the head than did raptors chasing aerial or aquatic prey and obligate scavengers. Ground predators that forage on mammals from above have a wide but short bill – which increases ingestion rate – and a large suborbital ridge to avoid sun glare. This may explain the protruding binocular field and the wide blind area above the head. By contrast, species from the two other groups have long but narrow bills used to pluck, flake or tear food and may need large visual coverage (and reduced suborbital ridges) to increase their foraging efficiency (e.g. using large visual coverage to follow the escaping prey in three dimensions or detect conspecifics). We propose that binocular field shape is associated with bill and suborbital ridge shape and, ultimately, foraging strategies. Highlighted Article: Raptors from Accipitriformes family differ in the shape of their binocular field according to their foraging tactics.

Evolutionary inference from QST-FST comparisons: disentangling local adaptation from altitudinal gradient selection in snapdragon plants

Phenotypic differentiation among natural populations can be explained by natural selection or by neutral processes such as drift. There are many examples in the literature where comparing the effects of these processes on multiple populations has allowed the detection of local adaptation. However, these studies rarely identify the agents of selection. Whether population adaptive divergence is caused by local features of the environment, or by the environmental demand emerging at a more global scale, for example along altitudinal gradients, is a question that remains poorly investigated. Here, we measured neutral genetic (FST) and quantitative genetic (QST) differentiation among 13 populations of snapdragon plants (Antirrhinum majus) in a common garden experiment. We found low but significant genetic differentiation at putatively neutral markers, which supports the hypothesis of either ongoing pervasive homogenisation via gene flow between diverged populations or reproductive isolation between disconnected populations. Our results also support the hypothesis of local adaptation involving phenological, morphological, reproductive and functional traits. They also showed that phenotypic differentiation increased with altitude for traits reflecting the reproduction and the phenology of plants, thereby confirming the role of such traits in their adaptation to environmental differences associated with altitude. Our approach allowed us to identify candidate traits for the adaptation to climate change in snapdragon plants. Our findings imply that environmental conditions changing with altitude, such as the climatic envelope, influenced the adaptation of multiple populations of snapdragon plants on the top of their adaptation to local environmental features. They also have implications for the study of adaptive evolution in structured populations because they highlight the need to disentangle the adaptation of plant populations to climate envelopes and altitude from the confounding effects of selective pressures acting specifically at the local scale of a population.

Genotype by environment interactions for the shade avoidance syndrome in the plant Antirrhinum majus, the snapdragon

A classical example of phenotypic plasticity in plants is the set of trait changes in response to shade, i.e. the shade avoidance syndrome. There is widespread evidence that plants in low light conditions often avoid shade by growing taller or by increasing their photosynthetic efficiency. This plastic response is expected to have evolved in response to selection in several species, yet there is limited evidence for its genetic variation within populations, which is required for any evolutionary response to selection. In this study, we investigated the shade avoidance syndrome in snapdragon plants (Antirrhinum majus) by using a common garden approach on four natural populations from the Mediterranean region. Our results showed that, in the four populations, individual plants reacted strongly to the presence of shade by growing longer shoots, longer internodes, and increasing their specific leaf area. Our results also revealed genetic variation for the plastic response within these populations, as well as few genetic constraints to its evolution. Our findings imply that the plastic response to shade has the potential to evolve in response to selection in natural populations of A. majus.

Olive tree varieties cultivated for the great Baetican oil trade between the 1st and the 4th centuries ad: morphometric analysis of olive stones from Las Delicias (Ecija, Province of Seville, Spain)

During the excavations of a Roman amphora workshop and oil mill of the 1st–4th century ad in Las Delicias, Genil valley, Ecija, Spain, large quantities of charred olive stones were recovered. The assemblages discovered in the pottery kilns demonstrate the use as fuel of olive residues, which were obtained from the extraction of the oil in the nearby mill. The abundance of material offered the opportunity to study the infra-specific diversity of the olives growing in the province of Baetica, which is known to have been an important oil-producing region during the Roman Empire. In total, 335 intact charred archaeological olive stones were analysed using geometric morphometry (outline analysis) and compared with several current morphotypes. These have been identified within a set of dimensional references of the stones established from the morphometric study of current varieties and wild populations, including genuinely wild and feral forms of olives, from various areas around the Mediterranean. The morphotype mainly found in wild populations was widely represented among the olive stones from Las Delicias. A large proportion of the archaeological stones were however close to various domesticated forms, which reflect the history of the region and of its varied cultural Mediterranean influences, Punic, Greek and Roman. Moreover, intermediate forms between two distinct morphotypes were identified. They suggest that hybrid olive trees derived from crosses among domesticated varieties and also between domesticated and wild forms, were grown in Las Delicias. In the Genil valley, Roman olive cultivation was based on a set of local olives which included wild and domesticated varieties from various origins, and whose diversity arose from breeding for improvement of varieties.