Limnological imbalances: an antipodean viewpoint

SUMMARY. 1. It is argued that modern limnology is excessively concerned with the study of fresh waters in the northern temperate region. Limnology has been and is unbalanced in its interests and emphases and, as a result, many widely held limnological concepts need revision. 2. The genesis of the argument is outlined. It involved the initial assumption that Australia was limnologically distinctive. Later, it was realized that since most of the world is arid, semi-arid, tropical or semi-tropical, Australia was much less distinctive than originally thought. 3. Other limnological assumptions considered in the paper concern saline lakes and salinization, the importance and nature of temporary bodies of standing fresh water, biological diversity in tropical fresh waters, the evolutionary importance of permanent freshwater lakes, and the global applicability of the River Continuum Concept. Consideration suggests that some widely-held views on these matters are erroneous. 4. Thus, saline lakes are more important (scientifically, commercially), widespread and numerous environments than is generally thought. Salinization is a significant environmental hazard for many rivers in semiarid regions. Temporary bodies of standing fresh water are geographically widespread and abundant aquatic environments with high species diversity, and in which many species have restricted distributions and low dispersal abilities. Biological diversity appears to be low in tropical fresh waters. Permanent freshwater lakes are perhaps less important loci for the evolution of the freshwater biota than generally thought, temporary fresh waters more so. Finally, the River Continuum does not seem to be applicable to many running waters outside the north temperate region. 5. A number of suggestions are made of how a better balance in limnology may be arrived at: how a more globally comprehensive intellectual framework may be built. Suggestions include: an increased awareness of the frequently different natures of waters within and beyond the northern temperate region; recognition of the potential benefits to be gained from the study of waters outside the northern temperate region;     

Adaptive patterns of osmotic and ionic regulation, and the invasion of fresh water by the palaemonid shrimps

To evaluate trends in the osmoregulatory behavior of neotropical, palaemonid shrimps, we investigated osmotic and ionic regulatory patterns in five species of Palaemon or Macrobrachium. The species' life histories depend on saline water to differing degrees, their habitats ranging from the marine/intertidal (P. northropi), through estuaries (P. pandaliformis) to coastal, freshwater streams (M. olfersii, M. potiuna) and inland, continental river systems (M. brasiliense). Hemolymph osmolality, chloride, sodium and magnesium concentrations were measured in shrimps exposed to experimental media ranging from fresh water (<0.5 per thousand ) to concentrated seawater (42 per thousand ) for up to 10 days. The marine and estuarine Palaemon species exhibit well-developed hyper/hypo-osmotic, sodium and chloride regulatory capabilities in mid-range salinities, tending to hyperconform in low salinities. The freshwater Macrobrachium species show variable hyperosmotic, sodium and chloride regulatory capacities, tending to hypoconform or unable to survive at higher salinities. All species hyper-regulate magnesium in fresh water, but hyporegulate strongly in saline media. Palaemonids from the saline habitats show the strongest osmoregulatory capabilities, and fresh water may have been gradually invaded by ancestral species with similar regulatory capacity. However, this regulatory plasticity has been lost to varying degrees in extant freshwater species.     

Fremde Fische in Flüssen und Seen

Non‐native fishes in rivers and seas The number of non‐native fish species in our water bodies is increasing in the course of globalisation. They navigate artificial canals, travel in ballast water of ships or are displaced as eggs attached to ships and other materials. Many of these species remain unnoticed but some may establish a reproductive population and even cause harm. Invasive gobies from the Black Sea – especially the bighead goby and round goby – are taken as an example to explain how biology and ecology can be studied and how we proceed to estimate the potential risk. Simple egg traps may help to reduce dispersal of these fish. Such measures are especially promising at locations such as the Upper Rhine where hydropower dams present bottlenecks.     

Invasion Genetics of Ponto-Caspian Gobies in the Great Lakes: A ‘Cryptic’ Species, Absence of Founder Effects, and Comparative Risk Analysis

Genetic variability and structure of nonindigenous vs native populations are compared for the Eurasian round goby Neogobius melanostomus and the tubenose goby Proterorhinus marmoratus, which both invaded Lake St. Clair of the North American Great Lakes about 1990. The round goby spread rapidly to all of the Great Lakes and the tubenose goby largely has been restricted to Lake St. Clair, with some recent range extension into western Lake Erie. Risk analyses may indicate whether genetic variability of colonizers is predictive of their relative invasive and establishment successes. The present investigation examined DNA sequence variation across the left domain of the mitochondrial DNA cytochrome b gene in round and tubenose gobies from Eurasian and Great Lakes locations. We also sequenced six additional Neogobius species (including the monkey N.␣fluviatilis, racer N. gymnotrachelus, and bigheadN. kessleri gobies that have been ‘on the move’ in Europe) and the knout goby Mesogobius batrachocephalus from the Black Sea in order to develop diagnostic genetic characters to identify them in case of future and/or undetected invasions and to delineate their phylogenetic relationships. Results show that a diverse number of haplotypes characterize round and tubenose goby populations from both North America and Eurasian sites, fitting a risk analysis prediction of high genetic variability in their successful introductions. Phylogenetic results indicate that the current genus Neogobius is paraphyletic and that the subgenusApollonia thus should be elevated to the level of genus, containingApollonia (N.) melanostomus (the round goby) andA. (N.) fluviatilis (the monkey goby). In addition, there appear to be two separate species of Proterorhinus marmoratus, a marine P. marmoratus Pallas 1814 in the Black Sea (matching the original type locality), and a ‘cryptic’ freshwater species in the Danube and Dnieper Rivers and probably other Eurasian freshwater habitats, as well as invasive in the Great Lakes. We suggest resurrecting the name P. semilunaris Heckel 1837 for the freshwater species (a taxon that was originally described from rivers draining into the Aegean Sea and the Danube River, but was later placed in synonymy with P. marmoratus). An erratum to this article is available at .     

Seasonal habitat use indicates that depth may mediate the potential for invasive round goby impacts in inland lakes

1. The round goby (Neogobius melanostomus) is among the fastest-spreading introduced aquatic species in North America and is radiating inland from the Great Lakes into freshwater ecosystems across the landscape. Predicting and managing the impacts of round gobies requires information on the factors influencing their distribution in habitats along the invasion front, yet this information is not available for many recently invaded ecosystems. We evaluated the seasonal habitat use and biomass of round gobies in an inland temperate lake to define the spatiotemporal scope of biological interactions at the leading edge of the round goby invasion.
2. Using novel statistical approaches, we combined hierarchical models that control for imperfect species detection with flexible smooth terms to describe non-linear relationships between round goby abundance and environmental gradients. Subsequently, we generated accurate detection-corrected estimates of the standing stock biomass of round gobies.
3. Our results show seasonally differentiated habitat niches, where suitable round goby habitat in summer months is restricted to shallow depths (<18.4 m) with a mixture of vegetative and mussel cover. We found high round goby biomass of 122 kg/ha in occupied habitats during the summer, with a total lake-wide biomass of 766,000 kg. In winter, round gobies migrate to deep offshore habitats and disperse, dramatically altering their scope for biological interactions with resident aquatic species across summer and winter seasons.
4. The results of this study indicate that the scope of biological interactions in inland lakes may be seasonally variable, with potential for high round goby biomass in shallow lakes or at the periphery of deep lakes in the summer months. Such shallow-water habitats may therefore present higher risk of ecological impacts from round gobies in invaded lentic ecosystems. As round gobies expand inland, consideration of seasonal habitat use will be an important factor in predicting the impacts of this pervasive invader.

     

Delivery of hatching larvae to estuaries by an amphidromous river shrimp: tests of hypotheses based on larval moulting and distribution

1.  Amphidromous shrimps live and breed in freshwater rivers and streams, but their larvae require development in sea water. Larvae may hatch upstream and then drift to the sea, although in some species females have been reported to migrate to the coast before larvae are released. Here, we tested the relative importance of larval drift and female migration in Macrobrachium ohione (Decapoda: Palaemonidae) in a distributary of the Mississippi River in Louisiana, U.S.A.
2.  Newly hatched (stage-1) larvae are nonfeeding and will not moult to stage 2 (first feeding stage) without encountering salt water. A factorial experiment was conducted in the laboratory to test the effects on moulting to stage 2 of (i) time spent by stage-1 larvae in fresh water before (ii) exposure to and maintenance in water of different salinity. Larvae kept in fresh water for 1 or 3 days before a change to saline water at 6 or 10 ppt showed a greater frequency of moulting than those kept for longer (5 days) in fresh water or changed to less saline water (2 ppt). Non-moulting larvae died or were moribund within 11 days of hatching.
3.  The relative abundance of stage-1 larvae was measured with plankton tows at two locations in the river c. 150 km apart, one near the sea and one upstream. Larval abundances near the sea were significantly greater than those upstream.
4.  The results indicate that hatched larvae of M. ohione have a limited period in which to drift in fresh water before reaching water sufficiently saline to stimulate moulting to the first feeding stage. Female migrations may play an important role in delivering larvae of amphidromous species from large continental river systems in which distances to the sea are great, while larval drift alone may be sufficient in species living in short streams, like those found in many small mountainous tropical islands.     

Redistribution of extracellular water and sodium may contribute to saline tolerance in wild ducks

The compartmentalization of body fluids was measured in three species of ducks that differ in saline tolerance. Half of the birds of each species drank freshwater, while the other half drank saline (300 mM NaCl). Among ducks that drank freshwater, total body water (TBW) was similar among all species, but Barrow's goldeneyes (Bucephala islandica), the most marine species, had larger extracellular fluid volume (ECFV) than freshwater mallards (Anas platyrhynchos) or estuarine canvasbacks (Aythya valisineria). When acclimated to saline, only goldeneyes shifted extracellular water and Na+ into the intracellular compartment. ECFV was correlated with plasma aldosterone concentration in goldeneyes, but not in canvasbacks (aldosterone was not measured in mallards). Data summarized from the literature showed that TBW does not differ among terrestrial, freshwater, or marine species, but marine species have a larger part of their TBW in the extracellular compartment. Saline induced movement of extracellular water and Na+ into the cells only in goldeneyes. ECFV and redistribution of extracellular water and Na+ into the cells may be important components in saline tolerance of marine birds.     

Molecular identification of two strains of third-stage larvae of Contracaecum rudolphii sensu lato (Nematoda: Anisakidae) from fish in Poland

Contracaecum sp. larvae (L3) from fish were identified using nucleotide sequences of the internal transcribed spacers ITS-1 and ITS-2 of the ribosomal DNA. The nematode larvae originated from fish in a freshwater situation (crucian carp Carassius carassius, from Selment Wielki Lake in Mazury, northeastern Poland) and a brackish-water region (Caspian round goby Neogobius melanostomus from the Baltic Sea, Gdafisk Bay at the Polish coast). Two strains (Contracaecum rudolphii A and B) of Contracaecum rudolphii senso lato, a parasite common at the adult stage in fish-eating birds, were identified. In fish from the freshwater site, only the strain temporarily designated C. rudolphii B was identified; in the brackish-water region, both strains were found, suggesting that fish serve as paratenic host for both genotypes. Contracaecum rudolphii sensu lato has been recorded in several species of fish-eating birds in Poland, particularly in the great cormorant, Phalacrocorax carbo, in which the abundance is highest. The results, although based on a restricted number of larvae, suggest that the life cycles of both genotypes can be completed in the Polish region and that at least one of them, C. rudolphii B, can develop both in fresh and brackish water.     

Secondary invasion of the round goby into high diversity Great Lakes tributaries and species at risk hotspots: potential new concerns for endangered freshwater species

The round goby (Neogobius melanostomus) first invaded North America in 1990 when it was discovered in the St. Clair River. Despite more than 15 years of potential invasion, many Great Lakes’ lotic systems remained uninvaded. Recently, we captured the round goby from several Great Lakes tributaries known as species-at-risk hotspots. With a combination of field sampling of round gobies and literature review of the impact of round gobies on native taxa, we assess the potential impacts of the secondary invasion to native species using three mechanisms: competition; predation; and indirect impacts from the loss of obligate mussel hosts. We estimate that 89% (17/19) of benthic fishes and 17% (6/36) of mussels that occur in these systems are either known or suspected to be impacted by the secondary invasion of round goby. In particular, we note that the distribution of potential impacts of round goby invasion was largely associated with species with a conservation designation, including seven endangered species (1 fish, 6 mussels). As these recent captures of round goby represent novel occurrences in high diversity watersheds, understanding the potential impacts of secondary invasion to native biota is fundamental to prevent species declines and to allow early mitigation.     

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     

Does the reproductive status of male round gobies (<Emphasis Type="Italic">Neogobius melanostomus</Emphasis>) influence their response to conspecific odours?

While pheromone communication has been well documented in the female round goby, male responses have not been examined. We determined if male round gobies responded to odours of their conspecifics, and whether that response varied with their reproductive status (reproductive and non-reproductive). No significant differences were observed for any treatment (dechlorinated water, or water conditioned by reproductive males (RM) and non-reproductive males (NRM) and females), although RM spent more time inside shelter than NRM (p < 0.05). Morphologically, length and weight did not vary, but gonadosomatic index, seminal vesicle somatic index and relative head width did differ between the two male groups (p = 0.000). Round goby males guard nests in cavities, and so a preference for shelter with increasing reproductive status is expected. Also, a lack of male responses to conspecific odours may be adaptive in this species, since males nest together in high densities, where increased aggression might prove detrimental for nesting males.     

Histological Studies on the Skin of a Freshwater Goby Rhinogobius brunneus (Gobiidae) Related to Cutaneous Respiration

Journal of Ichthyology - The Korean freshwater goby, Rhinogobius brunneus, prefers shallow water bodies with pebble bottoms, which are easily exposed to air due to the lack of water in droughts or...     

Spread of alien (non-indigenous) fish species <Emphasis Type="Italic">Neogobius melanostomus</Emphasis> in the Gulf of Gdansk (south Baltic)

The round goby (Neogobius melanostomus) was first noticed in the Gulf of Gdansk in 1990. This Ponto-Caspian fish was most likely introduced to the Baltic environment from ships ballast water. During the first years of invasion, slow population growth was observed in the initially colonized regions. From 1994, round goby gradually spread in all shallow water zones of the Gulf of Gdansk. The area occupied by the population and the number of fish grew rapidly. By the end of the nineties, round goby became a dominant fish in shallow waters of the western part of the Gulf of Gdansk, where the fish used all solid substrates on the bottom, including concrete piers, big stones or even dumped waste for spawning and refuge. The presence of round goby in the Vistula Lagoon and shallow waters near Rugia (Western Baltic) was noticed for the first time in 1999. The present state, size and condition of the round goby population in the Gulf of Gdansk led us to predict a continual spread of this species into new regions of the Baltic. In addition, the population of round goby is so large that the species has started playing a part in commercial fishing in the Gulf of Gdansk.     

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 .     

Standing genetic diversity and selection at functional gene loci are associated with differential invasion success in two non‐native fish species

Invasive species are expected to experience a unique combination of high genetic drift due to demographic factors while also experiencing strong selective pressures. The paradigm that reduced genetic diversity should limit the evolutionary potential of invasive species, and thus, their potential for range expansion has received little empirical support, possibly due to the choice of genetic markers. Our goal was to test for effects of genetic drift and selection at functional genetic markers as they relate to the invasion success of two paired invasive goby species, one widespread (successful) and one with limited range expansion (less successful). We genotyped fish using two marker types: single nucleotide polymorphisms (SNPs) in known‐function, protein‐coding genes and microsatellites to contrast the effects of neutral genetic processes. We identified reduced allelic variation in the invaded range for the less successful tubenose goby. SNPs putatively under selection were responsible for the observed differences in population structure between marker types for round goby (successful) but not tubenose goby (less successful). A higher proportion of functional loci experienced divergent selection for round goby, suggesting increased evolutionary potential in invaded ranges may be associated with round goby's greater invasion success. Genes involved in thermal tolerance were divergent for round goby populations but not tubenose goby, consistent with the hypothesis that invasion success for fish in temperate regions is influenced by capacity for thermal tolerance. Our results highlight the need to incorporate functional genetic markers in studies to better assess evolutionary potential for the improved conservation and management of species.     

On the finding of round goby <Emphasis Type="Italic">Neogobius melanostomus</Emphasis> (Pallas) (Gobiidae) in the food of burbot <Emphasis Type="Italic">Lota lota</Emphasis> (L.) of the Rybinskoe water reservoir

The fish swallowed by the burbot caught with gill nets in the Volzhskii reach of the Rybinskoe water reservoir in 2007 are investigated. Based on the teretial body shape, the absence of a swim bladder, the presence of a pelvic disc, the dimensions of the skull, the body length, the body weight, and the age, the specimens found in the gut of the burbot are identified as round goby Neogobius melanostomus (Pallas). The round goby should be regarded as an adventive (invasive) species because it is not distributed in different biotopes of the Rybinskoe water reservoir.     

Correlation between body size and fatty acid and essential amino acid composition of round goby (Neogobius melanostomus) and monkey goby (Neogobius fluviatilis) from the Rhine River (Germany)

In this study correlations between body size and muscle fatty and amino acid content of two species of goby, round goby (Neogobius melanostomus) and monkey goby (Neogobius fluviatilis) caught from river Rhine (Germany) were investigated. Among saturated fatty acids (SFAs), mono- (MUFA) and polyunsaturated fatty acids (PUFAs) only SFAs were significantly higher in round goby than monkey goby (P < 0.05). In general, the correlation between body size of both gobies and the content of most of the individual fatty acids was not significant. In monkey goby, the content of palmitic acid (C16:0) and oleic acid (C18:1 n-9) was positively correlated with weight (r = 0.43) and total length (r = ?0.58), respectively, and the content of docosahexaenoic acid (DHA) increased with condition factor (r = 0.50). The content of threonine, arginine, valine, phenylalanine and isoleucine in monkey goby was higher than those of round goby (P < 0.05). In round goby the three essential amino acids arginine, valine and leucine were positively (P < 0.05) correlated with body length, which indicates that longer round gobies are of higher nutritional value.     

Evolution of physiological tolerance and performance during freshwater invasions

Invasive species that penetrate habitat boundaries are likelyto experience strong selection and rapid evolution. This studydocuments evolutionary shifts in tolerance and performance followingthe invasion of fresh water by the predominantly estuarine andsalt marsh copepod Eurytemora affinis. Common-garden experimentswere performed on freshwater-invading (Lake Michigan) and ancestralsaline (St. Lawrence marsh) populations to measure shifts inadult survival (at 0, 5, and 25 PSU), and survival during developmentand development time (both using full-sib clutches split across0, 5, 15, and 25 PSU). Results showed clear evidence of heritableshifts in tolerance and performance associated with freshwaterinvasions. The freshwater population exhibited a gain in low-salinitytolerance and a reduction in high-salinity tolerance relativeto the saline population, suggesting tradeoffs. These tradeoffswere supported by negative genetic correlations between survivalat fresh (0 PSU) versus higher salinities. Mortality in responseto salinity occurred primarily before metamorphosis, suggestingthat selection in response to salinity had acted primarily onthe early life-history stages. The freshwater population exhibitedcurious patterns of life-history evolution across salinities,relative to the saline population, of retarded development tometamorphosis but accelerated development from metamorphosisto adulthood. This pattern might reflect tradeoffs between developmentrate and survival in fresh water at the early life-history stages,but some other selective force acting on later life-historystages. Significant effects of clutch (genotype) and clutch-by-salinityinteraction (G x E) on survival and development time in bothpopulations indicated ample genetic variation as substrate fornatural selection. Variation for high-salinity tolerance waspresent in the freshwater population despite negative geneticcorrelations between high- and low-salinity tolerance. Resultsimplicate the importance of natural selection and document theevolution of reaction norms during freshwater invasions.     

How Close Do We Live to Water? A Global Analysis of Population Distance to Freshwater Bodies

Traditionally, people have inhabited places with ready access to fresh water. Today, over 50% of the global population lives in urban areas, and water can be directed via tens of kilometres of pipelines. Still, however, a large part of the world''s population is directly dependent on access to natural freshwater sources. So how are inhabited places related to the location of freshwater bodies today? We present a high-resolution global analysis of how close present-day populations live to surface freshwater. We aim to increase the understanding of the relationship between inhabited places, distance to surface freshwater bodies, and climatic characteristics in different climate zones and administrative regions. Our results show that over 50% of the world''s population lives closer than 3 km to a surface freshwater body, and only 10% of the population lives further than 10 km away. There are, however, remarkable differences between administrative regions and climatic zones. Populations in Australia, Asia, and Europe live closest to water. Although populations in arid zones live furthest away from freshwater bodies in absolute terms, relatively speaking they live closest to water considering the limited number of freshwater bodies in those areas. Population distributions in arid zones show statistically significant relationships with a combination of climatic factors and distance to water, whilst in other zones there is no statistically significant relationship with distance to water. Global studies on development and climate adaptation can benefit from an improved understanding of these relationships between human populations and the distance to fresh water.