Stable isotope niche differentiation in sticklebacks with symmetric and asymmetric pectoral fins

Fluctuating asymmetry (FA) is often, but controversially, viewed as an indicator of fitness and a target of selection. In the brook stickleback, Culaea inconstans (Kirtland), FA of the pectoral fins, which are the main source of propulsion, is inversely correlated with fecundity. We examined the hypothesis that asymmetry of the pectoral fins could affect locomotion in such a way as to influence foraging and niche use in prereproductive brook stickleback. Nitrogen and carbon stable isotope analysis showed the diet of symmetric and asymmetric males diverged with increasing body size. Larger symmetric males fed at higher trophic levels and had a diet based on carbon emanating from a more pelagic source than their asymmetric counterparts. Such effects were not observed in females or smaller males. The number of chironomid larvae found in the gut was greater on average in asymmetric than symmetric fish. The results from this study strongly suggest FA of pectoral fins affects the foraging behaviour of C. inconstans and that stable isotope analyses of individual phenotypes provides a useful tool for assessing the ecological consequences of FA.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 617–623.     

Major morphological changes in a Lake Victoria cichlid fish within two decades

During the upsurge of the introduced predatory Nile perch in Lake Victoria in the 1980s, the zooplanktivorous Haplochromis ( Yssichromis ) pyrrhocephalus nearly vanished. The species recovered coincident with the intense fishing of Nile perch in the 1990s, when water clarity and dissolved oxygen levels had decreased dramatically due to increased eutrophication. In response to the hypoxic conditions, total gill surface in resurgent H. pyrrhocephalus increased by 64%. Remarkably, head length, eye length, and head volume decreased in size, whereas cheek depth increased. Reductions in eye size and depth of the rostral part of the musculus sternohyoideus, and reallocation of space between the opercular and suspensorial compartments of the head may have permitted accommodation of larger gills in a smaller head. By contrast, the musculus levator posterior, located dorsal to the gills, increased in depth. This probably reflects an adaptive response to the larger and tougher prey types in the diet of resurgent H. pyrrhocephalus . These striking morphological changes over a time span of only two decades could be the combined result of phenotypic plasticity and genetic change and may have fostered recovery of this species.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 41–52.     

Body size of the red fox Vulpes vulpes in Spain: the effect of agriculture

The body size of animals is affected by several factors, including ambient temperature and food availability. Ambient temperature is often negatively related to body size (Bergmann's rule) whereas an improved diet, especially during growth, has a positive effect. Animals commensal with man commonly exploit additional food sources (e.g. garbage dumps), thereby increasing their food supply. Using museum material, we studied morphological variation in skull size (and thus body size) among Spanish red foxes. Four measurements were taken of each skull and were related to the habitat from which the foxes were collected (agricultural and non-agricultural), and to latitude as a proxy for ambient temperature. The skull size of foxes collected in agricultural areas during the late 20th Century was significantly larger than that of those from non-agricultural areas, and was negatively related to latitude, thus contradicting Bergmann's rule. We suggest that increased food availability from animal husbandry is the cause for the observed increase in skull size (and thus body size).  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 729–734.     

Divergence in trophic ecology characterizes colonization of extreme habitats

Extreme habitats are characterized by the presence of physio‐chemical stressors, but also differ in aspects of the biotic environment, such as resource availability or the presence of competitors. The present study quantifies variation in trophic ecology of a small livebearing fish (Poecilia mexicana, Poeciliidae) across four different habitats that included nonsulphidic and sulphidic surface waters, as well as a nonsulphidic and a sulphidic cave. Resource use in different habitat types was investigated using gut content analysis. Populations diverged in resource use from a diet dominated by algae and detritus in nonsulfidic surface habitats to a diet including invertebrate food items in the other habitats. Poecilia mexicana in cave habitats further exhibited a higher dietary niche width than conspecifics from surface habitats. The condition of P. mexicana was analysed using storage lipid extractions. Fish from sulphidic and cave habitats exhibited a very poor condition, suggesting resource limitation and/or high costs of coping with extreme conditions. Finally, divergence in resource use was correlated with variation in viscerocranial morphology. A common garden experiment indicated both a genetic and plastic basis to the morphological variation observed among field populations. It is suggested that the morphological diversification is an adaptation to the differential use of resources among populations. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 517–528.     

Comparison of digestive efficiency in the parasitic great spotted cuckoo and its magpie host nestlings

Altricial nestlings are under strong selection pressures to optimize digestive efficiency because this is one of the main factors affecting nestling growth and survival. Bird species vary in their ability to assimilate different nutrients and current theory predicts that nestlings should also be able to adjust their nutritional physiology to feeding frequency. Variation in parental provisioning to nestlings would select for flexibility in nestling digestive physiology, which would allow maximization of nutrient assimilation. In the present study, by making use of a brood parasite–host study system in which great spotted cuckoo nestlings (Clamator glandarius) are reared by magpie (Pica pica) host foster parents when sharing the nest with host nestlings, we tested several predictions of the adaptive digestive efficiency paradigm. A hand‐feeding experiment was employed in which we fed both great spotted cuckoo and magpie nestlings with exactly the same diet simulating one food abundance period and one food deprivation period. The results obtained show that cuckoo nestlings ingested more food, gained significantly more weight during the abundance period, and assimilated a higher proportion of the ingested food than magpie nestlings. These results demonstrate for the first time that cuckoo nestlings enjoy digestive adaptations that favour a rapid processing of the ingested food, thereby maximizing their intake rate but without decreasing digestive efficiency. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 280–289.     

Dietary ratio of protein to carbohydrate induces plastic responses in the gastrointestinal tract of mice

Some vertebrates change the size of their digestive system in response to quantity and fibre content of ingested food, but the effects of dietary nutrients on gut structure remain poorly understood. Here we investigate how the protein to carbohydrate ratio of diets affects the mass of the gastrointestinal tract in mice. We fed 6-week-old male mice one of five isocaloric diets differing only in protein to carbohydrate ratio (the “no-choice” treatments), while a further four treatment groups received nutritionally complementary food pairings from which they could self-select a diet (the “choice” treatments). After 32 days, we measured the resulting dry mass of stomachs, intestines, caeca and colons. In the no-choice treatments, the stomachs were heavier in the mice fed diets containing more protein and less carbohydrate, indicating that larger stomachs may be needed for efficient digestion of the protein-rich food. In contrast, intestines, caeca and colons were heavier when diets contained more carbohydrates and less protein. This response may function to increase the digestive rate of carbohydrates when the dietary content of this macronutrient increases, but it may also indicate a compensatory response to increase amino acid uptake from a protein-deficient food. Mice in the choice treatments self-selected a diet with a protein to carbohydrate ratio of 0.46, and had gut dimensions similar to the expectation derived from no-choice treatments for this diet composition. Our results provide an example of plasticity in the differential allocation of resources to organ function, which is triggered by variation in resource quality.     

Incipient speciation driven by phenotypic plasticity? Evidence from sympatric populations of Arctic charr

Recent models suggest that the existence of environmentally induced polymorphisms within a single population (especially those related to foraging) facilitates the process of evolutionary divergence within a single gene pool by generating distinct phenotypic modes that are exposed to differential selection. In order to test a prediction of the phenotypic plasticity model of divergence, we used a well-documented polymorphism to disentangle the relative effects of morph and rearing environment in generating phenotypic variance. We reared first-generation offspring of two sympatric morphs of Arctic charr Salvelinus alpinus in the laboratory and compared their head morphology with that of their wild parents. Morphological characters with a known functional role in foraging were highly plastic. Rearing environment accounted for the largest component of the variation in expressed phenotype, but this environmental effect overlaid a clear (but small) genetic effect. We conclude that phenotypic plasticity has played a significant role in the evolution of this trophic polymorphism, but that the evolutionary process has progressed to the point that the gene pool is now segregated.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 611–618.     

Divergent walleye (Sander vitreus)-mediated inducible defenses in the centrarchid pumpkinseed sunfish (Lepomis gibbosus)

Predation has important ecological and evolutionary consequences. Evolutionary responses to diversifying selection include genetic differentiation, the evolution of adaptive phenotypic plasticity, and the genetic differentiation of plastic responses between populations. We tested if pumpkinseed sunfish ( Lepomis gibbosus ) respond to predation cues by changing their external body form in functionally sensible ways. We then asked whether predation has influenced the divergence of coexisting littoral and pelagic ecomorphs, by testing for divergent predator-induced responses. Juvenile L. gibbosus of both ecomorphs were reared with and without predation cues supplied by walleye ( Sander vitreus ) feeding on L. gibbosus . Predation cues stimulated increased body depth and dorsal spine length, but no increase in anal spine length or pectoral fin size. The dorsal spines of pelagic ecomorphs also grew longer than did those of littoral ecomorphs, while positive body depth responses were similar in both ecomorphs. This is the second fish taxa in which predator-induced morphological responses have been found, and the first in which divergent responses have been detected between ecomorphs. This suggests that the developmental systems of L. gibbosus ecomorphs have diverged under selection related to predation. We propose that other 'resource polymorphisms' in fishes have evolved under selection arising from a variety of factors, including predation, and not just selection related to resource use.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 25–36.     

Latitudinal and ecological correlates of morphological variation in Galaxias platei (Pisces, Galaxiidae) in Patagonia

We examined the morphological variability (23 morphometric traits) among individual Galaxias platei ( N  = 380) collected from 20 postglacial lakes in the southern Andes. The lakes were chosen to cover the latitudinal range of the species in Patagonia. Diet examined for a subset of these fish ( N  = 261) collected during the summer months (January through March), differed among river basins but there were no clear latitudinal trends in the consumption of any prey type. Diet may, however, have partially shaped morphology, as pelvic measures were negatively correlated with consumption of amphipods. Substantial differentiation among populations was observed, primarily in the shape and dimensions of the head, in caudal morphology and in fin length. Our results indicate that the morphology of G. platei varies with latitude, and may be related to risk of predation and diet.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 69–82.     

Differences in intestinal microbiota between avian brood parasites and their hosts

The intestinal microbiota determines the effectiveness of digestion in vertebrates, and is influenced by the external environment (mainly the diet), gut characteristics, and phylogeny. Avian brood-parasitic nestlings of the sub-family Cuculinae develop in nests of phylogenetically distant passerines and can be fed with the host diet. If the shaping of bacterial communities is dominated by phylogenetic constraints, and therefore the microbiota of parasitic nestlings differs from that of host nestlings, the energy and micronutrients that parasites and hosts obtain from a similar amount of food would be different. In this case, the bacterial communities of parasitic and host nestlings would have important consequences with respect to brood parasite development. By experimentally creating mixed broods of magpies ( Pica pica ) and great spotted cuckoos ( Clamator glandarius ), we investigated their cloacal microbiota using ribosomal intergenic spacer analysis. We found significant differences in bacterial assemblages of the parasitic and host nestlings, although none of the phylotypes were specific in either great spotted cuckoos or magpies. Cuckoos presented more complex communities, which could help the brood parasitic life style and allow the digestion of food provided by different potential hosts. Moreover, the intestinal morphology is different between the two species due to phylogenetic differences in the two taxa, which would influence the dissimilar bacterial assemblages. The detected differences in microbiota of great spotted cuckoo and magpie nestlings, which might occur in other brood parasite–host systems, may imply a lower digestion efficiency in parasites. Thus, the higher level requirements of cuckoo nestlings may be explained, at least in part, by cuckoos having a suboptimal bacterial community for processing the host diet.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 406–414.     

Edible wild plants of the Chorote Indians, Gran Chaco, Argentina

The Chorote Indians are hunter-gatherers and fishermen from north-west Argentina and south-west Paraguay who belong to the Mataco-Maká linguistic family. Their edible plants are identified by botanical and vernacular names, the parts employed and modes of preparation and consumption. The Chorote people use 57 plant species as a source of food, which they consume in 118 different ways. Five new edible species, that yield seven plant foods, are reported here for the first time for Chaquenian ethnic groups. However, only a few wild plant foods are in frequent use today, with most being used occasionally, infrequently or not at all. A cross-cultural comparison with four neighbouring ethnic groups reveals that one third of their plant foods are exclusive to the Chorote people, despite the fact that they share most of their edible plants with the other groups. This article is the first contribution to an understanding of the Chorote's ethnobotany and to a comparison of interethnic relationships concerning their edible plant resources.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 73–85.     

Genetic and morphological divergence along the littoral–pelagic axis in two common and sympatric fishes: perch,Perca fluviatilis (Percidae) and roach,Rutilus rutilus (Cyprinidae)

Individuals are constantly in competition with one another and, on both ecological and evolutionary timescales, processes act to reduce this competition and promote the gain of fitness advantages via diversification. Here we have investigated the genetic (AFLP) and morphological (geometric morphometrics) aspects of the littoral–pelagic axis, a commonly observed resource polymorphism in freshwater fishes of postglacial lakes. We found a large degree of variation in the genetic and morphological divergence between littoral and pelagic perch and roach across Swedish lakes. Although there was evidence of assortative mating (elevated kinship values) in both species, we could not find any significant coupling of morphology and genetic divergence. Instead, there was evidence that the extent of resource polymorphism may be largely caused by phenotypic plasticity. These results suggest that assortative mating, which can lead to genetically determined adaptive divergence, does occur in these species, particularly perch, but not according to genetically fixed morphological traits. The behavioural mechanisms facilitating associative mating need to be investigated to explore the interaction between phenotypic plasticity and adaptive genetic divergence and their roles in diversification. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 929–940.     

Trophic polymorphism in a riverine fish: morphological, dietary, and genetic analysis of mountain whitefish

Trophic polymorphisms are a prominent form of phenotypic diversification in many animal taxa. Northern temperate lakes have become model systems for the investigation of sympatric speciation due to trophic polymorphisms. Many examples of niche-based phenotypic variation occur in temperate lakes, whereas northern rivers offer few such examples. To further investigate the conditions under which trophic polymorphisms are likely to evolve, the present study examined phenotypic variation related to snout size and shape in the mountain whitefish (Salmonidae: Prosopium williamsoni ), which has been hypothesized to exhibit a rare example of reproductively isolated trophic morphs in a northern river-dwelling fish species. Variation in snout size and shape increased greatly with body size and, although this variation was continuously distributed, individuals in the largest size class tended to lie at phenotypic extremes. At one extreme were individuals with a large bulbous snout and a sloping forehead ('pinocchio'), and at the other were individuals that lack the bulbous snout and have a concave forehead ('normal'). The pinocchio trait may result from a stage-specific developmental switch that occurs late in ontogeny. Consistent differences were found with respect to diet between individuals with extreme snout morphologies, but no evidence was found for assortative mating within populations at seven microsatellite loci. The explosive mating system of this species may be responsible for this lack of assortative mating. The present study highlights the influence of ecological factors in shaping phenotypic and behavioural diversification due to trophic morphology.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 253–267.     

Size-related changes in cranial morphology affect diet in the catfish Clariallabes longicauda

Within the catfish family Clariidae, species exist with different degrees of jaw adductor hypertrophy. This jaw adductor hypertrophy has been related to bite performance, in turn suggesting a link to dietary specialization. Thus, an increase in the degree of hypertrophy will likely be reflected in an increase in the amount of hard prey in the diet. In the present study, we examine the ontogenetic scaling of cranial structure and diet in a species of catfish with a moderate degree of jaw adductor hypertrophy, Clariallabes longicauda . Additionally, we investigate whether the observed changes in the morphology of the feeding system during growth are linked to changes in diet. The fish examined demonstrate a strong positively allometric growth of the jaw adductors, of head height and of maximal head width, suggesting that larger fish can feed on larger and harder prey. Dietary data confirm these hypotheses and reveal an increase in maximal prey size consumed, the proportion of large prey in the diet, and average prey hardness during ontogeny. Moreover, the observed changes in the proportion of large prey consumed and prey hardness are correlated with an increase in lower jaw width and maximal head width, respectively. An increase in the amount of evasive prey in the diet with fish size is correlated with an increase in hyoid length. In summary, not only size dependent, but also size-independent variation of the feeding system was associated with ontogenetic changes in diet in C. longicauda .  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 323–334.     

An inquiline fig wasp using seeds as a resource for small male production: a potential first step for the evolution of new feeding habits?

The processes allowing evolutionary transitions in resources used by parasitic wasps are largely unknown. Microhymenopteran communities associated with figs could provide a model system to investigate such transitions. We investigate here a species of genus Idarnes . The larvae generally develop as inquilines of the pollinating wasp larvae. However, in figs where the parasitic pressure is high, eggs are laid in developing seeds. These eggs turn into small males. This is the first report of seed consumption by a fig wasp. Using an alternative resource to produce small males could provide a pathway to select for increased ability to use this resource and hence provide an intermediate step for evolving the capacity to use new resources.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 9–17.     

A new species of Ceratozamia (Zamiaceae) from the Sierra Madre of Chiapas, Mexico, with comments on species relationships

The new species Ceratozamia vovidesii from a Pleistocene floristic refuge in southern Mexico is described and illustrated. It show an affinity with C. matudae Lundell and C. mirandae Vovides, Pérez-Farrera & Iglesias from Chiapas, but it differs from them in leaf, male and female cones, and trunk morphology.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 393–400.     

Plastic resource polymorphism: effects of resource availability on Arctic char (Salvelinus alpinus) morphology

Resource polymorphism has been suggested to be a platform for speciation. In some cases resource polymorphism depends on phenotypic plasticity but in other cases on genetic differences between morphotypes, which in turn has been suggested to be the ongoing development of a species pair. Here we study environmentally induced morphological differences in two age classes of Arctic char ( Salvelinus alpinus ) influencing char performance and diet in relation to resource availability. We found that structurally complex habitats with relatively lower zooplankton densities gave rise to individuals with a deeper body, and a downward positioned tip of the snout compared with individuals from structurally simple habitats with relatively higher zooplankton densities for both age classes. Environment also had an effect on foraging efficiency on zooplankton, with fish from structurally simple habitats had a higher foraging rate than fish from structurally complex habitats. Diet analyses showed that resource use in char mainly depends on the relative abundance of different resources. Therefore, to gain further understanding of resource polymorphism we suggest that future studies must include population dynamic feedbacks by the resources on the consumers.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 85 , 341–351.     

Experimental evidence for adaptive phenotypic plasticity in a rock-dwelling cichlid fish from Lake Victoria

We demonstrate adaptive phenotypic plasticity (PP) in Neochromis greenwoodi , a rock-dwelling haplochromine cichlid from Lake Victoria. Adaptive phenotypic plasticity will enhance the chance of survival and reproduction in changing or novel environments. For rock-dwelling cichlids it may contribute to the success of settlement after migration between rocky outcrops or islands, since newly colonized patches of rock may offer different food regimes and different competitors. In our experiments, we simulated such situations by raising fry on different diets and with different competitors. We chose diet treatments in such a way that we could predict the direction of anatomical changes from functional demands. We expected an indirect effect of interspecific competition: selectively removing one prey-type by the competing species would leave N. greenwoodi with the other. We found the predicted phenotypic responses to food in trophic traits of N. greenwoodi . However, in the current experimental set-up we did not find a differentiating effect of species of competitor. We discuss the possible role of PP in allopatric and sympatric speciation.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 185−192.     

Leaf teeth in certain Salicaceae and 'Flacourtiaceae'

The leaf teeth glands in four taxa from Salicaceae and six from 'Flacourtiaceae' were examined using both light and scanning electron microscopes. There appears to be a progression from glands of simple structure in the flacourtiaceous taxa and a tendency to a more complicated development in morphology and anatomy of the salicaceous species.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 241–256.     

Contrasting Population and Diet Influences on Gut Length of an Omnivorous Tropical Fish,the Trinidadian Guppy (Poecilia reticulata)

Phenotypic plasticity is advantageous for organisms that live in variable environments. The digestive system is particularly plastic, responding to changes in diet. Gut length is the result of a trade-off between maximum nutrient absorption and minimum cost for its maintenance and it can be influenced by diet and by evolutionary history. We assessed variation in gut length of Trinidadian guppies (Poecilia reticulata) as a function of diet, season, ontogeny, and local adaptation. Populations of guppies adapted to different predation levels have evolved different life history traits and have different diets. We sampled guppies from sites with low (LP) and high predation (HP) pressure in the Aripo and Guanapo Rivers in Trinidad. We collected fish during both the dry and wet season and assessed their diet and gut length. During the dry season, guppies from HP sites fed mostly on invertebrates, while guppies in the LP sites fed mainly on detritus. During the wet season, the diet of LP and HP populations became very similar. We did not find strong evidence of an ontogenetic diet shift. Gut length was negatively correlated with the proportion of invertebrates in diet across fish from all sites, supporting the hypothesis that guppy digestive systems adapt in length to changes in diet. Population of origin also had an effect on gut length, as HP and LP fish maintained different gut lengths even in the wet season, when their diets were very similar and individuals in both types of populations fed mostly on detritus. Thus, both environment and population of origin influenced guppies gut length, but population of origin seemed to have a stronger effect. Our study also showed that, even in omnivorous fish, gut length adapted to different diets, being more evident when the magnitude of difference between animal and plant material in the diet was very large.