Bart J. A. Pollux

Native and non-native plants provide similar refuge to invertebrate prey, but less than artificial plants

Non-native species introductions are widespread and can affect ecosystem functioning by altering the structure of food webs. Invading plants often modify habitat structure, which may affect the suitability of vegetation as refuge and could thus impact predator-prey dynamics. Yet little is known about how the replacement of native by non-native vegetation affects predator-prey dynamics. We hypothesize that plant refuge provisioning depends on (1) the plant’s native status, (2) plant structural complexity and morphology, (3) predator identity, and (4) prey identity, as well as that (5) structurally similar living and artificial plants provide similar refuge. We used aquatic communities as a model system and compared the refuge provided by plants to macroinvertebrates (Daphnia pulex, Gammarus pulex and damselfly larvae) in three short-term laboratory predation experiments. Plant refuge provisioning differed between plant species, but was generally similar for native (Myriophyllum spicatum, Ceratophyllum demersum, Potamogeton perfoliatus) and non-native plants (Vallisneria spiralis, Myriophyllum heterophyllum, Cabomba caroliniana). However, plant refuge provisioning to macroinvertebrate prey depended primarily on predator (mirror carp: Cyprinus carpio carpio and dragonfly larvae: Anax imperator) and prey identity, while the effects of plant structural complexity were only minor. Contrary to living plants, artificial plant analogues did improve prey survival, particularly with increasing structural complexity and shoot density. As such, plant rigidity, which was high for artificial plants and one of the living plant species evaluated in this study (Ceratophyllum demersum), may interact with structural complexity to play a key role in refuge provisioning to specific prey (Gammarus pulex). Our results demonstrate that replacement of native by structurally similar non-native vegetation is unlikely to greatly affect predator-prey dynamics. We propose that modification of predator-prey interactions through plant invasions only occurs when invading plants radically differ in growth form, density and rigidity compared to native plants.

Consistent individual differences in seed disperser quality in a seed-eating fish

Animal-mediated seed dispersal (zoochory) is considered to be an important mechanism regulating biological processes at larger spatial scales. To date, intra-specific variation in seed disperser quality within seed-dispersing animals has not been studied. Here, I employed seed feeding trials to quantify individual differences in disperser quality within the common carp (Cyprinus carpio) using seeds of two aquatic plants: unbranched bur-reed (Sparganium emersum, Sparganiaceae) and arrowhead (Sagittaria sagittifolia, Alismataceae). I found substantial variation among carp individuals in their propensity to ingest seeds and their ability to digest them, resulting in up to 31-fold differences in the probability of seed dispersal. In addition, there were significant differences in the time that seeds are retained in their digestive systems, generating a twofold difference in the maximum distance over which they can potentially disperse seeds. I propose that seed-eating animal species consist of individuals that display continuous variation in disperser quality, with at one end of the continuum individuals that are likely to eat seeds, pass them unharmed through their digestive tract and transport them over large distances to new locations (i.e. high-quality seed dispersers) and at the other end individuals that rarely eat seeds, destroy most of the ones they ingest and transport the few surviving seeds over relatively short distances (low-quality seed dispersers). Although individual differences in seed dispersal quality could be the result of a variety of factors, these results underline the ecological and evolutionary potential of such variation for both plants and animals.

Locomotion during digestion changes current estimates of seed dispersal kernels by fish

Summary 1. Dispersal of seeds by animals is an important mechanism regulating plant diversity, range expansions and invasions. Many birds, mammals, fish and reptiles regularly ingest, transport and excrete viable seeds (known as endozoochory). 2. The effectiveness of endozoochory is modelled in dispersal kernels: functions that describe seed shadows in the landscape by combining movement of animals with experimentally obtained seed retention times and survival. 3. Currently, dispersal kernels use experimental data from resting animals, yet only moving animals disperse seeds. Although physical activity is known to affect digestive processes, little is known on how and to what extent this may influence current estimates of endozoochory. Activity may either prolong seed retention in the animal’s gut (locomotion-priority mode hypothesis) or may not affect seed excretion rate (digestion-priority mode hypothesis), and may affect seed survival and germination positively or negatively. 4. We tested how activity alters dispersal estimates in fish. We compared the seed dispersal potential of two riparian plant species (Carex acuta and Carex riparia) by the common carp (Cyprinus carpio) subjected to three different activity levels: low (basal metabolic rate, BMR), medium (2 9 BMR) or high activity (3 9 BMR). 5. Physical activity of the fish did not affect the number of intact retrieved seeds over 15 h of activity, but significantly affected seed retrieval patterns over time for both seed species. More active fish started seed excretion about 1 h later and kept excreting seeds at least 2 h longer. Effects of gut passage on germination could only be tested for C. acuta, where it reduced the percentage of germinating seeds by 22%, independent of the activity level. Seeds ingested by the fish germinated on average 3� 5d ays later than non-ingested control seeds. Seed retention times did not affect the timing of germination. 6. Our results support the locomotion-priority mode hypothesis and show that modelling dispersal kernels using parameters from inactive fish may underestimate potential dispersal distances. Because a trade-off between physical activity and digestive physiology is likely common in animals, it should be taken into account in future modelling of endozoochorous seed dispersal kernels.

Niche segregation in two closely related species of stickleback along a physiological axis: Explaining multidecadal changes in fish distribution from iron-induced respiratory impairment

Acute exposure to iron can be lethal to fish, but long-term sublethal impacts of iron require further study. Here we investigated whether the spatial and temporal distribution (1967–2004) of two closely related species of stickleback matched the spatial distribution of iron concentrations in the groundwater. We used the ‘Northern Peel region’, a historically iron-rich peat landscape in The Netherlands as a case study. This allowed us to test the hypothesis that niche segregation in two closely related species of stickleback occurred along a physiological axis. Patterns in stickleback occurrence were strongly associated with spatial patterns in iron concentrations before 1979: iron-rich grid cells were avoided by three-spined stickleback (Gasterosteus aculeatus, Linnaeus 1758) and preferred by nine-spined stickleback (Pungitius pungitius, [Linnaeus, 1758]). After 1979, the separation between both sticklebacks became weaker, corresponding to a decreased influence of local groundwater on stream water quality. The way both species changed their distribution in the field provides a strong indication that they differ in their susceptibility to iron-rich conditions. These observed differences correspond with differences in their respiration physiology, tolerance of poor oxygen conditions and overall life-history strategy documented in the literature. Our results exemplify how species can partition niche along a non-structural niche axis, such as sublethal iron-rich conditions. Other fish species may similarly segregate along concentration gradients in iron, while sublethal concentrations of other metals such as copper may similarly impact fish via respiratory impairment and reduced aerobic scope.

Acquired versus innate prey capturing skills in super-precocial live-bearing fish

Live-bearing fish start hunting for mobile prey within hours after birth, an example of extreme precociality. Because prenatal, in utero, development of this behaviour is constrained by the lack of free-swimming sensory-motor interactions, immediate success after birth depends on innate, evolutionarily acquired patterns. Optimal performance however requires flexible adjustment to an unpredictable environment. To distinguish innate from postnatally developing patterns we analysed over 2000 prey capture events for 28 metallic livebearers (Girardinus metallicus; Poeciliidae), during their first 3 days after birth. We show that the use of synchronous pectoral fin beats for final acceleration and ingestion is fixed and presumably innate. It allows for direct, symmetrical control of swimming speed and direction, while avoiding head yaw. Eye movements and body curvatures, however, change considerably in the first few days, showing that eye-tail coordination requires postnatal development. The results show how successful prey captures for newborn, live-bearing fish are based on a combination of fixed motor programmes and rapid, postnatal development.

Three-dimensional analysis of the fast-start escape response of the least killifish, Heterandria formosa

ABSTRACT Fish make C-starts to evade predator strikes. Double-bend (DB) C-starts consist of three stages: Stage 1, in which the fish rapidly bends into a C-shape; Stage 2, in which the fish bends in the opposite direction; and a variable Stage 3. In single-bend (SB) C-starts, the fish immediately straightens after Stage 1. Despite fish moving in three-dimensional (3D) space, fast-start responses of adult fish have mainly been studied in a horizontal plane. Using automated 3D tracking of multi-camera high-speed video sequences, we show that both SB and DB fast-starts by adult female least killifish (Heterandria formosa) often contain a significant vertical velocity component, and large changes in pitch (DB up to 43u2005deg) and roll (DB up to 77 deg) angles. Upwards and downwards elevation changes are correlated with changes in pitch angle of the head; movement in the horizontal plane is correlated with changes in yaw angle of the head. With respect to the stimulus, escape heading correlates with the elevation of the fish at the onset of motion. Irrespective of the initial orientation, fish can escape in any horizontal direction. In many cases, the centre of mass barely accelerates during Stage 1. However, it does accelerate in the final direction of the escape in other instances, indicating that Stage 1 can serve a propulsive role in addition to its preparatory role for Stage 2. Our findings highlight the importance of large-scale 3D analyses of fast-start manoeuvres of adult fish in uncovering the versatility of fish escape repertoire. Summary: The fast-start response of the least killifish entails a rich three-dimensional kinematic repertoire including complex combinations of roll, pitch and yaw motions.

Why do placentas evolve? Evidence for a morphological advantage during pregnancy in live-bearing fish

A live-bearing reproductive strategy can induce large morphological changes in the mother during pregnancy. The evolution of the placenta in swimming animals involves a shift in the timing of maternal provisioning from pre-fertilization (females supply their eggs with sufficient yolk reserves prior to fertilization) to post-fertilization (females provide all nutrients via a placenta during the pregnancy). It has been hypothesised that this shift, associated with the evolution of the placenta, should confer a morphological advantage to the females leading to a more slender body shape during the early stages of pregnancy. We tested this hypothesis by quantifying three-dimensional shape and volume changes during pregnancy and in full-grown virgin controls of two species within the live-bearing fish family Poeciliidae: Poeciliopsis gracilis (non-placental) and Poeciliopsis turneri (placental). We show that P. turneri is more slender than P. gracilis at the beginning of the interbrood interval and in virgins, and that these differences diminish towards the end of pregnancy. This study provides the first evidence for an adaptive morphological advantage of the placenta in live-bearing fish. A similar morphological benefit could drive the evolution of placentas in other live-bearing (swimming) animal lineages.