Personality-dependent inter- and intraspecific foraging competition in the invasive round goby,Neogobius melanostomus

This study examines the impact of boldness on foraging competition of the highly invasive round goby Neogobius melanostomus Pallas 1815. Individual risk tolerance, or boldness, was measured as the time to resume movement after a simulated predation strike. Fish that resumed movement faster were categorized as “bold,” fish that took more time to resume movement were categorized as “shy” and those that fell in between these two categories were determined to have “intermediate” boldness. Competitive impacts of boldness in N. melanostomus were determined in a laboratory foraging experiment in which interspecific (juvenile Atlantic cod Gadus morhua Linnaeus 1758) and intraspecific (intermediate N. melanostomus) individuals were exposed to either bold or shy N. melanostomus competitors. G. morhua consumed fewer prey when competing with bold N. melanostomus than when competing with shy N. melanostomus, whereas intermediately bold N. melanostomus foraging was not affected by competitor boldness. Bold and shy N. melanostomus consumed similar amounts of prey, and the number of interactions between paired fish did not vary depending on the personality of N. melanostomus individuals. Therefore, intraspecific foraging competition was not found to be personality dependent. This study provides evidence that individual differences in boldness can mediate competitive interactions in N. melanostomus; nonetheless, results also show that competition is also governed by other mechanisms that require further study.     

Temporal patterns in growth and survival of the round goby Neogobius melanostomus

Monthly, overwinter and annual instantaneous growth rates for round goby Neogobius melanostomus were calculated with maximal growth occurring in July and August and almost no growth observed between ice appearance (October) and melt (March). Annual absolute growth rates averaged 27·3 ± 1·9 mm for males and 19·8 ± 2·4 mm for females. The most parsimonious Cormack–Jolly–Seber model indicated that both the survival and recapture probabilities were dependent on sampling date, but not sex. Survival estimates remained high throughout the 13 month study with a median weekly survival probability of 0·920 (25 and 75% quartiles: 0·767 and 0·991), an overwinter survival probability of 99% and an annual survival rate of 67%. Survival probabilities were lowest for both sexes near the completion of the N. melanostomus reproductive season in July and August which supports existing evidence of higher mortality after reproduction, while challenging the paradigm that male N. melanostomus suffer comparatively higher mortality as a result of reproduction than females. Evidence indicating that growth and mortality rates are highest at the end of the reproductive season not only highlights seasonal variability in N. melanostomus natural history, but may also guide the control of this invasive species to periods when they are most vulnerable.     

Comparative feeding ecology of invasive Ponto-Caspian gobies

Invasions of Ponto-Caspian gobiid fishes are suspected to cause regime shifts in freshwater ecosystems. This study compared the trophic niche differentiations of Neogobius melanostomus and Ponticola kessleri in the upper Danube River using stable isotope analyses (δ¹³C and δ¹⁵N), gut content analyses and morphometric analyses of the digestive tract. Both species were identified as predacious omnivores with high dietary overlap and a generalistic feeding strategy. Amphipods (especially invasive Dikerogammarus spp.) contributed 2/3 to the index of food importance. δ¹⁵N-signatures of N. melanostomus revealed an ontogenetic diet shift and significantly exceeded those in P. kessleri by ~1.5‰, indicating a niche separation of half a trophic level. P. kessleri had shorter uncoiled intestinal tracts than N. melanostomus, indicating a narrower niche and adaptation to animal food. Trophic niches in both species expanded during the growth period with increasing intraguild predation and cannibalism in P. kessleri and increasing molluscivory in N. melanostomus. P. kessleri showed a higher degree of specialization and more stable feeding patterns across seasons, whereas N. melanostomus adapted its diet according to the natural prey availability. The feeding patterns of both species observed in the upper Danube River strongly differ from those in their native ranges, underlining their great plasticity. Both goby species consumed mainly other non-native species (~92% of gut contents) and seemed to benefit from previous invasions of prey species like Dikerogammarus villosus. The invasive success of gobies and their prey mirror fundamental ecological changes in large European freshwater ecosystems.     

Effects of sampling techniques on population assessment of invasive round goby Neogobius melanostomus

In this study, a comparison of point abundance sampling (PAS) electrofishing, angling with two different hook sizes and trap‐based fishing was performed in a non‐wadeable river to analyse their effects on catch per unit effort (CPUE) and population characteristics of invasive round goby Neogobius melanostomus. PAS electrofishing was identified as the most effective (mean ± s.e . CPUE = 57 ± 4 N. melanostomus min^?1) and least selective method in terms of size, feeding status and species composition. Angling had the second highest CPUE, but was more size selective and resulted in a higher proportion of males compared to electrofishing [overall sex ratio angling (female:male) = 1:0·92, electrofishing 1:0·65]. Owing to low CPUE (0·012 ± 0·004) and low frequency of occurrence, minnow traps were least suitable for N. melanostomus population assessment. The results of this study suggest that a higher degree of standardization and inter‐calibration is useful to achieve better comparability of population data of invasive N. melanostomus and other benthic fish species.     

What can morphology tell us about ecology of four invasive goby species?

This study presents a detailed comparative analysis of external morphology of four of the most invasive goby species in Europe (round goby Neogobius melanostomus, bighead goby Ponticola kessleri, monkey goby Neogobius fluviatilis and racer goby Ponticola gymnotrachelus) and interprets some ecological requirements of these species based on their morphological attributes. The results are evaluated within an ontogenetic context, and the morphological differences between the species are discussed in terms of the question: can special external shape adaptations help to assess the invasive potential of each species? The morphometric analyses demonstrate important differences between the four invasive gobies. Neogobius melanostomus appears to have the least specialized external morphology that may favour its invasive success: little specialization to habitat or diet means reduced restraints on overall ecological requirements. The other three species were found to possess some morphological specializations (P. kessleri to large prey, N. fluviatilis to sandy habitats and P. gymnotrachelus to macrophytes), but none of these gobies have managed to colonize such large areas or to reach such overall abundances as N. melanostomus.     

No effect of round goby <Emphasis Type="Italic">Neogobius melanostomus</Emphasis> colonisation on young-of-the-year fish density or microhabitat use

The round goby Neogobius melanostomus has recently invaded several major freshwater systems in Europe and North America. Despite numerous studies predicting an impact on native fish assemblages, few have attempted to demonstrate it. In this case study, we monitored the effect of N. melanostomus colonisation on abundance and habitat utilisation of both young-of-the-year (YOY) native fish and YOY western tubenose goby Proterorhinus semilunaris in a typical, medium-sized European river. Colonisation by N. melanostomus had no apparent effect on either native fish abundance and species richness or P. semilunaris abundance. Moreover, after colonisation, both native fish and P. semilunaris occupied similar niches (i.e. microhabitats) to those occupied before colonisation. While niche use of YOY N. melanostomus and P. semilunaris overlapped significantly, YOY native fish utilised different habitats from the gobiids. We suggest that N. melanostomus did not compete for resources with YOY fish in our study area due to lack of niche overlap and/or surplus resources. As N. melanostomus rapidly dominated the fish assemblage at our site, we further suggest that utilisation of an empty niche, rather than competitive superiority, was the main factor facilitating its success.     

荒漠草原土壤相对湿度对猪毛蒿表型可塑性的影响

猪毛蒿(Artemisia scoparia)为菊科蒿属草本植物,是一种适应性较强的广幅种。研究荒漠草原不同土壤相对湿度条件下猪毛蒿的表型可塑性,对认识异质生境下猪毛蒿的生存适应策略具有重要的生态学意义。结果表明:株高、茎粗、根长、根重和单株生物量均表现出随土壤相对湿度的增大而增加的趋势,对异质生境具有较强的可塑性,而根冠比则表现出相对的稳定性。植株不同部位生物量大小排序为:上部中部下部,且植株下部显著大于上部生物量(P0.05)。土壤相对湿度40%生境下的头状花序数量和重量显著高于土壤相对湿度30%和30%—40%生境。繁殖器官绝对投入量(lg R)随着个体大小(lg V)的增大呈极显著的增加(P0.001),繁殖阈值介于1.868—2.006 g。随着土壤相对湿度的增加,繁殖分配比例极显著增大(P0.001)。营养器官和繁殖器官生物量、头状花序重量和数量、地下生物量和地上生物量均呈极显著线性正相关关系(P0.001),存在正向权衡。单个头状花序重量并不随个体大小和头状花序数量的增加而发生显著变化(P0.05),且在不同土壤相对湿度和不同部位间均无显著差异(P0.05)。由此可见,猪毛蒿在异质生境下产生的可塑性是其生存繁殖的重要反应机制之一。     

Invasive round goby Neogobius melanostomus has sex‐dependent locomotor activity and is under‐represented in catches from passive fishing gear compared with seine catches

The higher proportion of males of the invasive round goby Neogobius melanostomus in samples from two activity selective passive fishing gears compared with one activity non‐selective fishing gear in three Dutch lakes is related to higher male locomotory activity and is a sex‐dependent trait. This difference in activity reflects the different ecology of male and female N. melanostomus.     

Evidence for sex‐specific selection in brain: a case study of the nine‐spined stickleback

Theory predicts that the sex making greater investments into reproductive behaviours demands higher cognitive ability, and as a consequence, larger brains or brain parts. Further, the resulting sexual dimorphism can differ between populations adapted to different environments, or among individuals developing under different environmental conditions. In the nine‐spine stickleback (Pungitius pungitius), males perform nest building, courtship, territory defence and parental care, whereas females perform mate choice and produce eggs. Also, predation‐adapted marine and competition‐adapted pond populations have diverged in a series of ecologically relevant traits, including the level of phenotypic plasticity. Here, we studied sexual dimorphism in brain size and architecture in nine‐spined stickleback from marine and pond populations reared in a factorial experiment with predation and food treatments in a common garden experiment. Males had relatively larger brains, larger telencephala, cerebella and hypothalami (6–16% divergence) than females, irrespective of habitat. Females tended to have larger bulbi olfactorii than males (13%) in the high food treatment, whereas no such difference was found in the low food treatment. The strong sexual dimorphism in brain architecture implies that the different reproductive allocation strategies (behaviour vs. egg production) select for different investments into the costly brains between males and females. The lack of habitat dependence in brain sexual dimorphism suggests that the sex‐specific selection forces on brains differ only negligibly between habitats. Although significance of the observed sex‐specific brain plasticity in the size of bulbus olfactorius remains unclear, it demonstrates the potential for sex‐specific neural plasticity.     

Plasticity of female reproductive resource allocation depends on the presence or absence of prior environmental sex determination in Ceratopteris richardii

Resource allocation plasticity enables individuals to alter patterns of nutrient use between reproductive and vegetative output to better fit their current environment. In sexually labile plant species, abiotic environmental factors can influence expression of dimorphic gender, resulting in environmental sex determination (ESD), which potentially reduces the need for plasticity of resource allocation by preemptively matching an individual’s future nutrient demands to resource availability in its location. Ceratopteris richardii gametophytes exhibit gender‐dependent differences in relative carbon and nitrogen content, and ESD in certain nutrient environments. This study examined whether prior ESD in C. richardii gametophyte populations reduced subsequent plasticity of reproductive allocation compared to instances where no ESD occurred, by quantifying phenotypic responses to reduced P, N, or CO₂ availabilities. All three nutrient‐limited environments resulted in decreased size of egg‐bearing (meristic) gametophytes compared to nonlimited environments, but gametophytes failed to respond to N and CO₂ limitation at the time of sex determination, resulting in no ESD. N limitation resulted in a predictable allometric re‐allocation of resources based on small gametophyte size, whereas CO₂ limitation caused a change in reproductive output consistent with true plasticity. Withholding exogenous P caused ESD and had no effect on relative reproductive output of resultant meristic gametophytes because the size decrease was minor. Under P limitation, ESD matched the resource demands of gender phenotypes to their environment before the onset of developmental dimorphism, reducing the need for large allocation adjustments after sex determination.     

Carcass feeding as a cryptic foraging mode in round goby Neogobius melanostomus

Round gobies Neogobius melanostomus were observed readily consuming soft tissue from carcasses of larger fishes under both laboratory and field conditions. Consumption normally progressed in a typical sequence, starting with soft and easily accessible tissues such as the eyes, followed by puncture of the abdominal cavity, gut consumption and then muscle consumption. Carcass feeding has not previously been seen in N. melanostomus and has potential consequences for transfer of nutrients and contaminants.     

Canalization of body size matters for lifetime reproductive success of male predatory mites (Acari: Phytoseiidae)

The adaptive canalization hypothesis predicts that highly fitness‐relevant traits are canalized via past selection, resulting in low phenotypic plasticity and high robustness to environmental stress. Accordingly, we hypothesized that the level of phenotypic plasticity of male body size of the predatory mites Phytoseiulus persimilis (low plasticity) and Neoseiulus californicus (high plasticity) reflects the effects of body size variation on fitness, especially male lifetime reproductive success (LRS). We first generated small and standard‐sized males of P. persimilis and N. californicus by rearing them to adulthood under limited and ample prey supply, respectively. Then, adult small and standard‐sized males were provided with surplus virgin females throughout life to assess their mating and reproductive traits. Small male body size did not affect male longevity or the number of fertilized females but reduced male LRS of P. persimilis but not N. californicus. Proximately, the lower LRS of small than standard‐sized P. persimilis males correlated with shorter mating durations, probably decreasing the amount of transferred sperm. Ultimately, we suggest that male body size is more strongly canalized in P. persimilis than N. californicus because deviation from standard body size has larger detrimental fitness effects in P. persimilis than N. californicus. © 2014 The Authors. Biological Journal of the Linnean Society published by John Wiley & Sons Ltd on behalf of The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 889–899.     

A comparison of phenotypic plasticity between two species occupying different positions in a successional sequence

We compared the phenotypic plasticity of two greenhouse-grown species (Corispermum macrocarpum and Salsola collina) occupying different positions in a successional sequence in Horqin Sandy Land, by treating with different population density and the availability of soil nutrients and water. The same species can exhibit different patterns of plasticity in response to different environmental factors. In the soil nutrient treatments, the plasticity pattern of S. collina could be described as “master-of-some”. However, in the soil-water and population-density treatments, it showed no significant difference from C. macrocarpum in the reaction norm for plasticity. It was similar to a “jack-of-all-trades” plasticity pattern. Contrary to the previous conclusion that late successional species had higher reproductive allocation than early successional species, in this successional sequence, the late species had lower reproductive allocation in all treatments. Reproductive allocation of both species increased with the increase in water availability and also increased with a decrease in nutrient levels. However, density had no effect on reproductive allocation. Although the root:shoot ratio increased with decreasing water availability, there were no differences in the plasticity pattern for this trait in both species. Root:shoot ratio was, however, not significantly affected by nutrient availability and density. In a word, the plasticity patterns of invaders are adapted to the analyses of succession.     

The sampling efficiency of electrofishing for Neogobius species in a riprap habitat: a field experiment

Even though electrofishing is commonly used to sample Neogobius spp. and other swimbladder‐lacking benthic fishes, its efficiency is considered poor especially in habitats with abundant interstitial spaces. To determine the efficiency of electrofishing Neogobius spp. and quantitatively estimate sampling bias in a riprap (shot rock used to armor shorelines against water erosion) mesohabitat, riprap fragments were set up in a natural riverine environment. The experimental setting enabled us to collect all fish remaining in the riprap fragments after these areas had been electrofished. The sampling efficiency of electrofishing Neogobius spp. (dominated by Neogobius melanostomus) varied between 17.6 and 47.4% (mean 29.7%), while percids (possessing a well‐developed swim bladder) were collected with 74.6% efficiency. Fish size had no effect on the probability of capturing Neogobius spp. by electrofishing. Within Neogobius spp., N. melanostomus was less susceptible to electrofishing than Neogobius gymnotrachelus (23.7% and 50.1%, respectively). Decreased electrofishing efficiency in areas of rocky substrate should be considered in estimates of total abundance of Neogobius spp., especially if they are to be compared with catches of other species possessing swim bladders.     

Intracanopy plasticity under strong wind conditions in the wild olive tree (Olea europaea L.): a conserved response between closely related taxa?

Trees have the ability to respond to local environmental cues by expressing particular phenotypes across their canopy through a mechanism known as intracanopy plasticity. In this study, intracanopy plasticity of Olea europaea subsp. europaea was analyzed by sampling leeward and windward canopy exposures of individuals occurring in an area with sustained strong wind conditions. A suite of morphofunctional and reproductive traits was measured at these contrasting canopy positions and, for comparison, also in wind-protected trees. Furthermore, the pattern of intracanopy plasticity of these plants was compared to that previously documented in a closely related species, Olea europaea subsp. guanchica. Plants exposed to strong winds displayed substantial differences between leeward and windward exposures in most of the study traits. Leeward exposures experienced a mean reduction of 73% in wind speed as compared to windward ones, and displayed a modular phenotype matching that observed in wind-protected plants. Wind-exposed plants, however, were comparatively smaller and had fewer and smaller inflorescences, since inflorescence size was positively associated with crown size. The two closely related species showed similar crown and leaf sizes between populations exposed to strong winds, and intracanopy responses were comparable for most traits. These observations suggest that intracanopy plasticity resulted in the expression of contrasting phenotypes within individuals, which allowed trees to persist under sustained wind stress, although at the cost of a reduced reproductive fitness. In addition, this study gives support to the idea that intracanopy responses are conserved among closely related taxa evolving in different habitats, but experiencing a comparable limiting factor.     

Phenotypic differentiation of Ponto-Caspian gobies during a contemporary invasion of the upper Danube River

Evolution is known to act on contemporary timescales and invasive organisms are often used to study rapid evolutionary changes of geno- and phenotypes under natural conditions. The ability and speed of local adaptation is discussed as a key character triggering successful invasions. Variation of body shape among populations of two highly invasive, sympatric Ponto-Caspian goby species (Teleostei: Gobiidae) with a comparable invasion history in the upper Danube River, i.e. Neogobius melanostomus and Ponticola kessleri, was assessed using geometric morphometric methods. Phenotypic variation established within less than 15 generations was evident in both species. It was mainly correlated with geographical location, but in N. melanostomus also with substrate type, an ecological variable reflecting habitat quality. The two species differed in their degree of intraspecific variation which was more pronounced in N. melanostomus, the numerically dominant invader in the upper Danube. Body shape variation significantly correlating with geographical rather than ecological variables suggests a heritable component and renders phenotypic plasticity as a lone explanation unlikely. Patterns of body shape similarity among upper Danubian goby populations suggest a stepping-stone rather than a continuous expansion model for both species, where multiple introductions, possibly from various origins, may have shaped differentiation.     

Getting What Is Served? Feeding Ecology Influencing Parasite-Host Interactions in Invasive Round Goby Neogobius melanostomus

Freshwater ecosystems are increasingly impacted by alien invasive species which have the potential to alter various ecological interactions like predator-prey and host-parasite relationships. Here, we simultaneously examined predator-prey interactions and parasitization patterns of the highly invasive round goby (Neogobius melanostomus) in the rivers Rhine and Main in Germany. A total of 350 N. melanostomus were sampled between June and October 2011. Gut content analysis revealed a broad prey spectrum, partly reflecting temporal and local differences in prey availability. For the major food type (amphipods), species compositions were determined. Amphipod fauna consisted entirely of non-native species and was dominated by Dikerogammarus villosus in the Main and Echinogammarus trichiatus in the Rhine. However, the availability of amphipod species in the field did not reflect their relative abundance in gut contents of N. melanostomus. Only two metazoan parasites, the nematode Raphidascaris acus and the acanthocephalan Pomphorhynchus sp., were isolated from N. melanostomus in all months, whereas unionid glochidia were only detected in June and October in fish from the Main. To analyse infection pathways, we examined 17,356 amphipods and found Pomphorhynchus sp. larvae only in D. villosus in the river Rhine at a prevalence of 0.15%. Dikerogammarus villosus represented the most important amphipod prey for N. melanostomus in both rivers but parasite intensities differed between rivers, suggesting that final hosts (large predatory fishes) may influence host-parasite dynamics of N. melanostomus in its introduced range.     

Geographic variation in phenotypic plasticity in response to dissolved oxygen in an African cichlid fish

Genetic adaptation and phenotypic plasticity are two ways in which organisms can adapt to local environmental conditions. We examined genetic and plastic variation in gill and brain size among swamp (low oxygen; hypoxic) and river (normal oxygen; normoxic) populations of an African cichlid fish, Pseudocrenilabrus multicolor victoriae. Larger gills and smaller brains should be advantageous when oxygen is low, and we hypothesized that the relative contribution of local genetic adaptation vs. phenotypic plasticity should be related to potential for dispersal between environments (because of gene flow’s constraint on local genetic adaptation). We conducted a laboratory‐rearing experiment, with broods from multiple populations raised under high‐oxygen and low‐oxygen conditions. We found that most of the variation in gill size was because of plasticity. However, both plastic and genetic effects on brain mass were detected, as were genetic effects on brain mass plasticity. F₁ offspring from populations with the highest potential for dispersal between environments had characteristically smaller and more plastic brains. This phenotypic pattern might be adaptive in the face of gene flow, if smaller brains and increased plasticity confer higher average fitness across environment types.     

Isolation and characterization of microsatellites in Neogobius kessleri (Perciformes,Gobiidae) and cross‐species amplification within the family Gobiidae

Fourteen polymorphic microsatellites were isolated from Neogobius kessleri, a benthic fish of Ponto‐Caspian origin which has been recently introduced into the Middle and Upper Danube River. Number of alleles and heterozygosity per locus in a sample of 32 fish individuals ranged from two to four and from 0.13 to 0.75, respectively. These primers will be useful in determining the population structure of N. kessleri. In addition, successful cross‐amplification was obtained for four related species, N. melanostomus, N. fluviatilis, N. gymnotrachelus and Proterorhinus marmoratus. These microsatellite loci may be useful for the evaluation of the origin of non‐native goby populations.