Life-history traits of invasive bighead goby Neogobius kessleri (Günther, 1861) from the middle Danube River, with a reflection on which goby species may win the competition

Life history traits of an invasive population of bighead goby Neogobius kesslerei (Günther, 1861) from the middle Danube, including absolute and relative fecundity, egg size, number of spawning batches and size at first maturation, were examined and evaluated within an epigenetic context. Ripe bighead goby females attained 42.8–142.5 mm L _S, with absolute fecundities ranging from 669 to 5646 eggs (mean 2109 eggs), and relative fecundities of 61.6–174.0 eggs g⁻¹ body weight (mean 119.6 eggs). Egg diameters varied between 0.04 mm and 1.70 mm (mean = 0.57 mm). In the pre-spawning period there was no clear size distinction in eggs (0.12–1.45 mm; mean = 0.52 mm) in 34.1% of females; whereas in 65.9% of females, two egg size groups were distinguished: group I diameters of 0.06–0.85 mm (mean = 0.43 mm), and group II diameters of 0.55–1.70 mm (mean = 1.17 mm). Females with size-group II eggs at the beginning of the reproductive season were assumed to be ready to spawn and the others to be subsequent spawners. Bighead goby appears to be altricial compared to the round goby, although in both species a shift from highly precocial towards a less precocial life history was observed. These differences, affected by epigenetic mechanisms and resulting in alternative ontogenies, may have important implications for a species' potential success in novel environments, favouring the round goby over short time periods (several years) and bighead goby over longer periods of time (decades and longer).     

Annual variations in the reproductive traits of Pomatoschistus microps in a Mediterranean lagoon undergoing environmental changes: evidence of phenotypic plasticity

The spawning period of the common goby Pomatoschistus microps from 1993 to 1997 in the Vaccarès lagoon did not vary, except in 1997 when it was longer due to the reproduction of the young-of-the-year. Egg size and number, and reproductive allocation varied greatly with one year to another. Female common gobies increased both their fecundity per spawning act and their egg size from 1993 to 1995. The annual variation in the reproductive effort suggests a high phenotypic plasticity of reproductive traits in P. microps , in the face of environmental perturbations. In winter 1993–1994, a centennial flood of the Rhône River caused major hydrological changes in the lagoon in less than 1 week, affecting many invertebrates and fish for several years. The reproductive investment of the common goby increased, possibly as a consequence of those environmental changes.     

Annual changes in brain concentration of arginine vasotocin and isotocin correspond with phases of reproductive cycle in round goby,Neogobius melanostomus

Reproductive cycle of seasonally breeding fish is synchronized with changes of photoperiod and temperature in environment. We hypothesize that arginine vasotocin (AVT) and isotocin (IT) are involved in timing and synchronization of seasonal reproductive activity in the round goby (Neogobius melanostomus). To verify this hypothesis, we examined the annual profiles of brain AVT and IT in round goby males and females in relation to their reproductive cycle. Wild round gobies were exposed to annual environmental changes in their natural habitats from where they were sampled monthly over a year. AVT and IT were measured using HPLC with fluorescence detection preceded by solid-phase extraction. This study shows seasonal variations in brain AVT and IT levels. Profiles of changes were similar in males and females: the peak of AVT was observed before spawning in March-April, whereas that of IT during spawning in May–June. Furthermore, the lowest AVT level was noted out of breeding season from November to January, while the level of IT decreased immediately at the end of the spawning. The results show that high AVT levels correlate with pre-spawning period whereas the highest IT levels correspond to spawning. A significant decline in AVT and IT in non-spawning season coincided with the quiescent phase of gametogenesis in both sexes.     

Ontogenetic variability in the external morphology of monkey goby, Neogobius fluviatilis (Pallas, 1814) and its relevance to invasion potential

In the previous decade, four species of non-native gobies have invaded the middle section of the river Danube and its tributaries. An effective tool for understanding biological invasions is the evaluation of various biological traits (morphological, life history, ontogenetic) within an epigenetic context. The present study examines the external morphology of monkey goby Neogobius fluviatilis (Pallas, 1814) from the mouth of the River Hron, the morphological differences among three goby species (monkey, bighead and round) and the relevance of these differences for invasive potential. Monkey goby reach their definite phenotype very early in their ontogeny and thus represent a strongly precocial (specialized) species with direct development. The morphological differences between monkey and two other goby species also reflect its strong specialization for sandy substrata and smaller prey types. Thus, monkey goby are not expected to spread to new areas as fast as the round and bighead gobies, and their distribution is likely to be limited to sandy and/or sandy-gravel substrata. If this assumption is correct, then the potential adverse impact of monkey goby on native fauna or even ecosystem is likely to be less than that of the bighead and round gobies. Handling editor: K. Martens     

Reproductive biology of racer goby Neogobius gymnotrachelus in the Włocławski Reservoir (Vistula River, Poland)

Initial studies on the reproductive parameters and spawning behaviour of racer goby Neogobius gymnotrachelus, one of three Ponto‐Caspian gobiid species recently invading Polish waters, were carried out in W?oc?awski Reservoir (lower River Vistula, Poland), where spawning lasted from April to August. Males were on average about 23.3% longer and 53.8% heavier than females. The smallest female with mature gonads was 58 mm TL (estimated age 2+ years), and males were sexually mature at 59 mm TL (age of 1+ years). Gonado‐somatic index at the beginning of spawning was seven times larger in females (mean = 8.4) than that in males (mean = 1.2). Mean absolute fecundity was 952 eggs per female (361–2236). Egg diameter ranged from 0.140 to 1.558 mm, being of bi‐modal distribution, which suggests two or three spawning bouts. Length–frequency analysis of juveniles revealed a multimodal distribution. Specialized reproductive behaviour (nest construction and parental care of eggs) and an extended spawning period increase the probability that racer goby can successfully establish self‐sustaining populations in novel environments.     

Little association of biological trait values with environmental variables in invasive alien round goby (Neogobius melanostomus)

The relative importance of species‐specific biological trait characteristics and environmental factors in invasions of nonindigenous species remains controversial because both have mostly been studied independently. Thus, the main objective of this study was to examine the correlation of biological traits with environmental variation in the globally invasive round goby Neogobius melanostomus from the upper Danube River. Based on a sample of 653 specimens along a continuous 200 km river pathway, links between nine environmental factors (substrate‐type, six water measurements, and the communities of fishes and macroinvertebrates) and seven biological traits (nutritional and energetic status, trade‐offs of parasite resistance and resource allocation, and three growth proxies) were analyzed. Biological trait values of N. melanostomus hardly correlated with the environment, could not explain invasion progress and imply a general low overall importance for invasion success. Instead, alternative individual life‐history trajectories appear to determine invasion success. This is in line with up to 15% of all specimens having outlying biological trait values of potential adaptive value, suggesting a considerable importance of adaptive trait variation among single individuals for the whole invasion progress. This “individual trait utility hypothesis” gives an alternative explanation for success of invasive species by single individuals carrying particular traits, and it should be specifically targeted and analyzed at currently invaded sites.     

Persistence of an invasive fish (Neogobius melanostomus) in a contaminated ecosystem

Post-establishment dynamics of invasive species have been under-studied. However, understanding these dynamics is particularly important for the management of invasive species known to impact native communities. Following the invasion of a highly invasive species, the round goby (Neogobius melanostomus), we document long-term population changes after establishment and address how population dynamics of a successful invader change through persistence and integration. Round goby present a threat to the areas they invade by out-competing native species for resources. Furthermore, as a pollution tolerant species, round goby present a second threat by acting as a possible vector for contaminant transfer to higher trophic levels in invaded ecosystems with areas of contamination. We sampled round goby for 11 years (2002–2012) at four low contamination sites and two high contamination sites within Hamilton Harbour ON, Canada, an International Joint Commission Area of Concern. Across sampling years, we show that round goby abundance has declined at low contamination sites, while remaining stable at high contamination sites. Moreover, we show that average body size decreased and reproductive investment increased both across sampling years and between sites of low and high contamination. Our results document population demographic shifts in a persisting invasive species, and underscore the importance of management practices for this species in contaminated environments.     

Evidence of a Male Sex Pheromone in the Round Goby (Neogobius melanostomus)

The reproductive success of the round goby (Neogobius melanostomus), an invasive fish, may be mediated by the use of pheromones. We hypothesized that reproductive male (RM) round gobies release sex pheromone to which reproductive females (RF) respond. In this study, we compared behavioural and electrophysiological responses of reproductive and non-reproductive female round gobies to conspeci fic males. Results of behavioural experiments in the laboratory showed that RF spent significantly more time near the source of the male odour compared with odours from control water. However, RF did not distinguish between odours from non-reproductive male (non-RM) water and control water. Non-reproductive females (non-RF) were not attracted to odours released from RM or non-RM water. Results of electro-olfactogram (EOG) responses showed that both RF and non-RF discriminated between HPLC fractionated RM and non-RM odours. However, the EOG responses of RF were about eight-fold higher than non-RF exposed to RM odours. These findings confirm that RM round gobies release a pheromone signal that attracts RF. The results of this research may be useful in developing control strategy using natural pheromones to disrupt the reproductive behaviour of the invasive round goby and to curtail its effects on native species. An erratum to this article is available at .     

Ontogenetic variability in external morphology of round goby Neogobius melanostomus from Middle Danube, Slovakia

In the previous decade, four species of non‐native goby have invaded the middle section of the River Danube. To understand the problems associated with these invasions better, we examined the external morphology of 184 round goby Neogobius melanostomus specimens using both triple regression (distance‐based measurements) and geometrical (coordinates‐based measurements) analysis within an ontogenetical and epigenetical aspect. Using geometrical analysis of distance‐based characters, we found that the external body shape in four size groups of round goby remained essentially unchanged. Such direct development represents a strongly precocial (i.e. specialized) life‐history, whereas earlier maturation in this non‐native population suggests a shift back towards more altricial (i.e. less specialized, more generalized) life history. This combination of altricial–precocial trajectories seems to be typical for invasive round gobies and may be one of key factors for their successful colonization of new environments.     

Age and reproductive investment in grass goby females in the Venice lagoon

The present work describes the relationship between age and reproductive investment in the grass goby females in the Venice Lagoon. Age was estimated by otoliths reading, while reproductive investment was assessed by either the relative number of mature females across the breeding season and their gonadosomatic index. Females from different size/age classes differed in the timing and level of reproductive investment. The oldest females (3+ years old) appeared earlier in the spawning habitat, investing more at the beginning of the breeding season. In turn a later arrival in the spawning habitats has been observed for younger females (0+) and smaller individuals of 1+ and 2+ age classes. Present results gave evidence of an earlier age/size at sexual maturity of the grass goby in the Venice Lagoon compared to previous information on the species from the same lagoon and from other Mediterranean areas. Results were discussed in the light of previous data on reproductive strategy of nesting males and of the relationship between fishing pressure and life history traits in the Venice lagoon.     

Yellow belly as honest signal of female quality in Knipowitschia panizzae(Gobiidae)

Sexually dimorphic traits are common in fish species, and examples from both males and females have been described. The function of these traits has been widely investigated in males. On the contrary, female ornaments have been studied mainly in sex role reversed species, such as pipefish, while their role in species with 'conventional' sex roles remain to be investigated.
This study focused on the presence, function, and possible role as indicator of female quality of a sexually dimorphic nuptial trait in the lagoon goby, Knipowitschia panizzae . In this species, that present conventional sex roles, females show a yellow spot on the belly. Aquarium spawning experiments demonstrated that the coloration on the belly is due to dermal pigments, is displayed only when female is ready to spawn and is switched off within few minutes from the end of egg deposition. This sexual trait presents variability in size among females and indicates female fecundity relatively to her own body size. As a consequence, female yellow belly appears to be an honest signal of female quality. Field data on natural nests highlighted that males perform parental cares mostly only on one egg batch at a time and the modality of egg deposition suggested that males are limited in their potential reproductive rates by environmental factors. Male limitation in egg care could constitute the basis for a female biased operational sex ratio, favouring male choosiness and the evolution of female nuptial displays.     

From sneaker to parental male: change of reproductive traits in the black goby, Gobius niger (Teleostei, Gobiidae)

This study focuses on the consequences of the switch of tactic from parasitic to parental male in the black goby, Gobius niger (Teleostei: Gobiidae), a species showing two alternative male mating tactics. Older and larger males defend nests, court, and perform parental care on eggs, while younger and smaller ones behave as parasites, sneaking into nests while spawning occurs. Males adopting different tactics are known to present differences in primary and secondary sex traits. The social context of sneaker males was manipulated to induce a tactic switch. Sneakers were kept under two different experimental treatments with or without a female, and under exclusion of male-male competition. Males changed tactics, courting females, spawning, and performing parental care. All males showed substantial changes in primary sexual traits, such as a reduction in gonadal development and an increase in the investment in accessory structures. The experimental groups differed in the functionality of gonads and accessory organs and in the development of the secondary sex traits. These results demonstrate that the moment of switching is not genetically fixed in the black goby. Sneaker males are able to quickly reallocate energy in primary and secondary sex traits, in accordance with the adopted tactic. Several aspects of this flexible reproductive pattern resemble the socially controlled sex change found in sequential hermaphrodites.     

Responses of round goby (<Emphasis Type="Italic">Neogobius melanostomus</Emphasis>) olfactory epithelium to steroids released by reproductive males

The wild perciform teleost Neogobius melanostomus (the round goby) originated from the Ponto-Caspian region and is now a highly successful invasive species in the Laurentian Great Lakes. Males may attract females into their nests for spawning by releasing reproductive pheromones, and it has been previously shown that reproductive males synthesize and release the 5β-reduced and 3α-hydroxyl steroids 3α-hydroxy-5β-androstane-11,17-dione (11-oxo-etiocholanolone; 11-O-ETIO) and 3α-hydroxy-5β-androstane-11,17-dione 3-sulfate (11-oxo-etiocholanolone-3-sulfate; 11-O-ETIO-3-s) and 3α,17β-dihydroxy-5β-androstan-11-one 17-sulfate. In this study, we investigated properties of these released steroids by recording field potential responses from the olfactory epithelium (electro-olfactogram, EOG). The steroid 3α,17β-dihydroxy-5β-androstan-11-one 17-sulfate did not elicit olfactory responses while both 11-O-ETIO and 11-O-ETIO-3-s stimulated olfactory field potentials in the round goby, but not in the goldfish. Cross-adaptation analysis demonstrated that round gobies discriminated between11-O-ETIO and 11-O-ETIO-3-s (as well as etiocholanolone, ETIO) at the sensory level. Second messenger cascades depending on both cAMP and IP₃ were inferred for steroids from pharmacological inhibition studies, while the canonical teleost odors taurocholic acid (a bile acid) and l-alanine (an amino acid) used only cAMP and IP₃, respectively. The round goby presents itself as an excellent species for the study of olfactory function of fish in the wild, given its possible use of these released steroids as pheromones.     

Food Resource Partitioning between Two Invasive Gobiid Species (Pisces,Gobiidae) in the Littoral Zone of the River Danube,Hungary

At present, the invasive round and bighead gobies are the most abundant benthic fish species in the near shore zone of the Middle Danube. We compared their diet seasonally in natural and artificial habitats and contrasted it with the food supply. The composition of the macroinvertebrate community was determined mainly by seasonal changes, whereas habitat type had smaller effect. Round gobies followed these changes flexibly. They consumed mainly chironomids in the spring, whilst amphipods and molluscs in the summer and autumn. Bighead gobies relied on amphipods in each season and in both habitats, and consumed fish, too, including round goby (intraguild predation). Diet overlap was determined by the morphological differences of the species allowing a varying degree of differentiation according to the seasonally changing food supply. (© 2009 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Reproductive biology of the freshwater goby Knipowitschia croatica Mrakovčić, Kerovec,Mišetić & Schneider 1996 (Actinopterygii,Gobiidae)

The reproductive biology and gonadogenesis cycle of the Vrgorac goby, Knipowitschia croatica is described [Conservation of Endangered Freshwater Fish in Europe, Birkhauser Verlag, Basel]. The species displays sexual dimorphism during the spawning period. Sexual maturity is achieved at an early age, with 50% of males and females sexually mature at total lengths of 40–45 mm. Fecundity of gravid females ranged from 188 to 593 eggs, with an egg diameter of 0.22–1.11 mm. Though the extended spawning period lasts from March to November, the highest intensity is observed from April to September. A comparison is made of the reproductive biology of this species with other sand goby species of the genera Knipowitschia and Pomatoschistus.     

Life‐history traits of temperate and thermophilic barracudas (Teleostei: Sphyraenidae) in the context of sea warming in the Mediterranean Sea

This study indicated that the life‐history traits of European barracuda Sphyraena sphyraena are apparently better suited to their environmental conditions compared to the more physically restricted life‐history traits of the yellow‐mouth barracuda Sphyraena viridensis, which co‐habit the north‐western Mediterranean Sea. The latter thermophilic species has a considerably higher reproductive potential as it invests its energy reserves in larger numbers of hydrated eggs per spawning batch. This would favour its population growth rates within the study area, especially if sea warming continues, in which case it is likely that the spawning phenology of this species would give it an advantage.     

Sublethal effects on reproduction in native fauna: are females more vulnerable to biological invasion?

Although invasive species are a major threat to survivorship of native species, we know little about their sublethal effects. In soft-sediment marine systems, mat-forming invasive species often have positive effects, facilitating recruitment and enhancing the diversity and abundance of native invertebrates. However, because mat-forming invasive species change the habitat in which they invade, and benthic invertebrates are sensitive to environmental disturbance, important sublethal effects on native species may exist. Using a model marine system we show that the widespread mat-forming invasive alga Caulerpa taxifolia (Vahl) C. Agardh has strong negative effects on the reproductive traits of a native bivalve Anadara trapezia (Deshayes, 1840) (e.g. timing of reproductive development and spawning, and follicle and gamete production) even though the invader has positive effects on recruitment. Moreover, gender specific responses occurred and indicated that females were more susceptible to invasion than males. Our results indicate that sublethal effects of an invasive species on reproductive traits will have severe consequences for fitness of the native species.     

Growth and reproduction of the non‐native icefish Neosalanx taihuensis Chen, 1956 (Salangidae) in a plateau lake,southwestern China

Growth, reproduction and abundance traits of the invasive icefish Neosalanx taihuensis Chen, 1956 were investigated monthly from July 2009 to May 2011 in Lake Erhai on the Yunnan‐Guizhou Plateau, south‐western China, in order to explore the changes in life‐history traits after translocation. The results indicated that the icefish exhibited obvious plasticity in growth and reproduction traits. Growth of the fish in Lake Erhai was faster than that in native waters and in other translocated reservoirs. By fitting the von Bertalanffy growth model to the data, it was estimated that icefish obtain an asymptotic size of 96.12 mm, a K of 1.61, and a t₀ of ‐0.26; the calculated overall growth performance index φ′ was 4.17. The strategy of reproduction changed from multiple‐ to single‐spawning. The spawning period was from October to December with the absolute and relative fecundities of 1250 ± 169 eggs per ind and 2557 ± 245 eggs per g, respectively. Plasticity in icefish growth and reproduction in Lake Erhai greatly facilitated its population establishment, making it one of the most abundant fish species. The icefish invasion in the lake may be one of the reasons for the decrease or extinction of native fish species populations, and some measures for the control of this invasive fish are suggested.     

Parasite fauna of native and non‐native populations of Neogobius melanostomus (Pallas, 1814) (Gobiidae) in the longitudinal profile of the Danube River

Parasite fauna of round goby Neogobius melanostomus (Pallas, 1814) in the Danube River was investigated in both its native range (two sites in the Bulgarian stretch of the Danube) and non‐native range of distribution (Croatian, Slovak and Austrian stretches) during 2005 and 2006. The aim was to identify possible changes in parasite communities associated with the introduction of a host into the new environment. A total of 29 metazoan parasite species were found to parasitize round goby in the Danube River; twelve of these parasite species were found in both the native and non‐native range of distribution. Introduction of a novel parasite species to the non‐native range via the round goby was not found. Eight parasite species occurred only in the native range and nine species only in the non‐native range of the round goby distribution. Losses of native parasite species in non‐native round goby populations and/or acquiring of novel parasite species in a new environment were not significant. Thirteen parasite taxa were recorded for the first time in round gobies. Three parasite taxa (Diplostomum spp., Pomphorhynchus laevis and Raphidascaris acus) were found in high prevalence and abundance at each sampling site in both the native and non‐native range. Parasite species diversity was assessed for each sampling site and season using three diversity indices (the Shannon, Simpson and Equitability indices), with the highest same‐season values found in a non‐native site in Slovakia (1.38, 0.69 and 0.60, respectively) and the lowest in a native site in Bulgaria (0.28, 0.12 and 0.14, respectively). Species diversity was higher in both non‐native round goby populations (Slovak and Austrian) compared to native Bulgarian populations. However, diversity indices values varied among almost all sampling sites.     

Finding round goby Neogobius melanostomus (Pallas, 1814) in the Rybinsk Reservoir

The data from finding a female round goby in June 2010 near the dam in the Central Reach of the Rybinsk Reservoir are presented. The presence of mature reproductive products may indicate the possibility of the successful acclimatization of this species in the Rybinsk Reservoir.