Asymmetrical Resource Ownership Increases Owners’ and Reduces Non‐Owners’ Motivation to Fight in the Mangrove Rivulus,Kryptolebias marmoratus

Resource ownership often increases an individual's aggressiveness and its probability of defeating a competitor. Individuals contesting resource owners could therefore incur higher costs, making individuals reluctant to compete with owners. We tested the hypothesis that animals use asymmetry in resource ownership as a cue for contest costs and adjust contest decisions accordingly. Using a mangrove rivulus fish (Kryptolebias marmoratus), we staged (1) contests with a randomly assigned asymmetry in resource ownership (one fish was provided with a shelter) and (2) contests in which neither fish had a shelter. Owners that were in their shelters at the contest start showed a greater tendency to fight and won more contests than their intruder opponents; those outside the shelter at the start did not. Compared with fish in contests with no shelters at stake, shelter owners had a higher tendency to fight whether or not they were in their shelters at the start; intruders, however, had a lower tendency to fight only against owners that were inside the shelter at the start. These results demonstrate (1) that ownership status influences both owners’ and intruders’ contest decisions (and in opposite directions), producing a detectable ownership advantage and (2) that intruders required confirmation of their opponents’ ownership status before retreating without challenging them. Ownership status per se is therefore important to the fish's contest decisions.     

Agonistic behaviour and control of access to hiding places in two intertidal blennies, Lipophrys pholis and Coryphoblennius galerita (Pisces: Blenniidae)

In this study we investigated the relationships between dominance rank and access to shelters in captive groups of Lipophrys pholis and Coryphoblennius galerita, as well as the effects of group size and shelter availability. Dominance rank was strongly correlated with size in juvenile L. pholis and with sex and size in adult C. galerita, males being dominant over females of similar size. Access to shelters was significantly correlated with dominance rank. For both species, most interactions occurred out of shelters. Direct disputes over shelters were always initiated by the dominant fish and the initiator was always the winner. The rate of aggression per fish per unit time decreased with an increase in the number of fish in L. pholis but not in C. galerita. No significant differences were found in groups differing in the number of shelters. C. galerita showed a higher rate of agonistic interactions and a higher proportion of overt aggression than L. pholis. It is suggested that one of the functions of agonistic interactions in these fishes is the control of a set of shelters, in the network of pathways used by each individual within its home range, minimising the time required to hide in case of danger. Received in revised form: 29 January 2001 Electronic Publication     

Occurrence of non‐native fishes in the Danube east of Vienna (Austria) and potential interactions of invasive gobiids with native fishes

Non‐native fish species pose a major threat to local fish populations and aquatic ecosystems in general. Invasive gobies are a particular focus of research, but with partly inconsistent results. While some studies reported severe detrimental impacts on native species, others have concluded less serious or neutral effects. We provide results from a large‐scale, multi‐annual fish monitoring program on the occurrence and abundance of non‐native fishes in the main stem of a free‐flowing section of the Austrian Danube. Special emphasis was placed on identifying positive or negative interactions of invasive gobies with native species. Whereas most non‐native species occurred too sporadically or were too few in number to infer a direct threat on the local fish community, invasive gobies were among the most common fishes throughout all sampling years. Co‐occurrence analyses revealed species‐ and mesohabitat type‐specific associations of gobies with native species, which were primarily positive. Notably, native predators such as asp, burbot, or perch probably benefit from the ubiquitous gobies. Two characteristic fluvial fishes revealed negative associations with invasive gobies, namely barbel (Barbus barbus) and Danube whitefin gudgeon (Romanogobio vladykovi): they appear to avoid habitats occupied by gobies. Accordingly, high abundances of round and bighead goby most likely resulted in population losses of barbel and whitefin gudgeon, respectively. Overall, our results indicate a limited negative impact of non‐native species in the sampling area. This is because only two out of 51 occurring species were found to be adversely affected by gobies, the share of co‐occurrences with native species was high, and other non‐native species were generally rare. Nevertheless, invasions are highly dynamic, and new non‐native species are likely to occur in the Austrian Danube, calling for continued monitoring and awareness.     

Vibrational Communication in the Cherry Leaf Roller Caterpillar Caloptilia serotinella (Gracillarioidea: Gracillariidae)

The cherry leaf roller (Caloptilia serotinella) produces three distinct types of substrate-borne signals—scraping, plucking, and vibrating—during interactions between conspecifics. Signals were recorded using a piezoelectric sensor, and behavioral experiments tested the hypothesis that signaling functions in territorial disputes over costly leaf shelters. Trials involving the introduction of a conspecific to a resident's leaf shelter demonstrated a significant increase in the amount of scraping by the resident; there was no significant difference in plucking or vibrating. In control trials, general mechanical disturbances such as opening and probing the shelter typically did not elicit signaling. Although both residents and intruders were observed to produce all three signal types, residents most often initiated signaling, and scraped significantly more than intruders. Implications for understanding the diversity of vibrational communication in larval Lepidoptera, particularly shelter-building species, are discussed.     

Male and female cooperatively breeding fish provide support for the "Challenge Hypothesis"

The idea that territorial aggression is regulated by androgensand that aggression itself can modulate androgen levels is wellestablished in males. In many species, females also displayaggressive behavior, yet little work has been conducted on theeffects of female aggression on hormone levels. In this study,we compared the effects of a simulated territory intrusion (amethod for testing the Challenge Hypothesis) on males and femalesof the fish, Neolamprologus pulcher. This cichlid fish fromLake Tanganyika is a particularly useful species to examinesex differences in the behavioral mediation of hormones as breedingpairs remain in a territory year round and both sexes defendthis territory against conspecific and heterospecific intruders.In our study, both sexes indeed aggressively defended theirterritory against a simulated territory intruder. In responseto intruders, both males and females displayed elevated levelsof circulating 11-ketotestosterone, but only females exhibitedincreases in testosterone. Neither aggressing male nor femalefish showed changes in estradiol levels compared to control(nonaggressing) fish. Residents were more aggressive than theintruders and won most of the interactions. However, residents(or winners) did not show higher hormone levels than intruders(or losers). We suggest that aggression commonly modulates androgenlevels in both male and female teleost fish.     

Experimental evidence that ecological effects of an invasive fish are reduced at high densities

Understanding the relationship between invasive species density and ecological impact is a pressing topic in ecology, with implications for environmental management and policy. Although it is widely assumed that invasive species impact will increase with density, theory suggests interspecific competition may diminish at high densities due to increased intraspecific interactions. To test this theory, we experimentally examined intra- and interspecific interactions between a globally invasive fish, round goby (Neogobius melanostomus), and three native species at different round goby densities in a tributary of the Laurentian Great Lakes. Eighteen 2.25 m² enclosures were stocked with native fish species at natural abundances, while round gobies were stocked at three different densities: 0 m^?2, 2.7 m^?2, and 10.7 m^?2. After 52 days, native fish growth rate was significantly reduced in the low density goby treatment, while growth in the high density goby treatment mirrored the goby-free treatment for two of three native species. Invertebrate density and gut content weight of native fishes did not differ among treatments. Conversely, gut content weight and growth of round gobies were lower in the high goby density treatment, suggesting interactions between round gobies and native fishes are mediated by interference competition amongst gobies. Our experiment provides evidence that invasive species effects may diminish at high densities, possibly due to increased intraspecific interactions. This is consistent with some ecological theory, and cautions against the assumption that invasive species at moderate densities have low impact.     

Associations between food habits and pharyngeal morphology in the round goby (<Emphasis Type="Italic">Neogobius melanostomus</Emphasis>)

Several studies have shown that round gobies (Neogobius melanostomus) undergo a dietary shift from arthropods to dreissenid mussels as they grow, and this shift is accompanied by changes in pharyngeal morphology associated with durophagy. In contrast, some populations of round gobies prey heavily on various arthropods, but it is unknown whether those populations develop pharyngeal morphology associated with durophagy or if they develop less robust and molarized structures. To test if there is a relationship between food habits and pharyngeal morphology, we compared those characters in round gobies from two sites in Erie Co., Pennsylvania: a dreissenid-present site (Presque Isle Bay, PIB, of Lake Erie) and a dreissenid-absent site (Fairview Gravel Pit, GP). Multivariate analysis of covariance (MANCOVA) revealed consistent ontogenetic changes at both sites in which lower pharyngeals of larger fish were more robust, possessed wider teeth, and had a greater area taken up by large diameter teeth. Pharyngeal morphology also differed between sites, with gobies from PIB having wider and more robust lower pharyngeals. Food habits differed markedly between round gobies from the two sites, with dreissenid mussels being the most important prey item for all length classes in PIB, and crustaceans being the most important prey item for all length classes at the GP. Canonical correlation analysis on all round gobies revealed a correlation between consumption of dreissenid prey and pharyngeal characters associated with durophagy. Although food habits and pharyngeal morphology of round gobies appear to be associated, the mechanism responsible for the association (i.e. phenotypic plasticity versus local adaptation) is not clear.     

Matter of Size or Matter of Residency Experience? Territorial Contest in a Green Hairstreak,Chrysozephyrus smaragdinus (Lepidoptera: Lycaenidae)

Territorial residents usually win asymmetrical owner‐intruder contests and a variety of hypotheses have been proposed to explain this phenomenon. In the butterfly Chrysozephyrus smaragdinus (Lepidoptera: Lycaenidae), male territorial residents defended their territories against intruders during numerous contests and kept them for many successive days. Field observations and experiments were conducted to examine the factors related to this superiority of residents. Forewing length did not differ between residents and intruders, suggesting that body size is not correlated with resource holding potential. Removal–replacement experiments demonstrated that residency did not serve as an arbitrary means for contest settlement, and did not support the recently presented alternative hypothesis that males with higher body temperature are more likely to win. New residents fought longer in defense of the territory as their residence duration in the territory increased. I discuss these results in light of game theory and suggest that the superiority of residents in C. smaragdinus may be based on the asymmetry of resource (territory) value for residents and intruders.     

Predation of zebra mussels by round gobies, Neogobius melanostomus

Preliminary gut analysis of a recent Great Lakes invader, the round goby, Neogobius melanostomus (7.0–8.4 cm), collected from the Detroit River, showed that they ate zebra mussels (58%), snails (6%), and other invertebrates (36%), including aquatic insects (Hexagenia), softshelled crayfish, and zooplankton. Because zebra mussels, Dreissena polymorpha, predominated as prey, we investigated the ability of round gobies to consume different size classes of zebra mussels. In laboratory experiments, we examined feeding preferences of three size classes of round gobies (5.5–6.9 cm; 7.0–8.4 cm; 8.5–10.3 cm standard length) on four different size classes of zebra mussels (6.0–9.9 mm, 10.0–12.9 mm, 13.0–15.9 mm, 16.0–18.9 mm). All sizes of round gobies ate zebra mussels < 10.0 mm. Only the largest size class of round gobies ate larger zebra mussels (10.0–12.9 mm) when all prey sizes were presented. The association between the total mass of zebra mussels available and the amount consumed by round gobies increased positively up to about 6.5 g of available mussels and then levelled off. Round gobies consumed an average of 1.0 g of mussels in 24 h. There was a significant positive relationship between gape size and standard length of round gobies. Although larger round gobies (over the size range of fish in our study) are able to consume larger zebra mussels, small mussels were preferred. Our findings suggest that the preference of small zebra mussels by round gobies has the potential to alter the size structure of zebra mussel populations. This revised version was published online in July 2006 with corrections to the Cover Date.     

Has the invasive round goby caused new links in Baltic food webs?

The Ponto-Caspian round goby (Neogobius melanostomus, Pallas 1814) most probably was established in the Gulf of Gdańsk, Baltic Sea, in the late 1980’s and has since become one of the dominant species in the region. In this study we assess the role of round gobies as prey for two important fish species in the Gulf of Gdańsk, cod (Gadus morhua) and perch (Perca fluviatilis). We compared their present diet with stomach analyses from the area prior the round goby establishment, as well as with diet analysis from Baltic regions where round gobies are absent. There were large differences in the diet between cods from the Gulf of Gdańsk 2003–2006 compared to cods in earlier studies (1977–1981) from the Southern Baltic Sea. There were also large differences in cod and perch diets from areas with and without round goby. Presently, round goby constitutes the most important prey for medium sized cods in Gulf of Gdańsk, and perch from the same area almost exclusively feed on gobiids. Stomach analysis, trophic level estimates, and stable isotope analyses all indicated that cod and perch in Gulf of Gdańsk after the round goby establishment belonged to a similar trophic level. Beside round goby, no mussel feeding fish contributed much to the diet of cod or at all to the diet of perch. Thus, it is likely that round gobies constitute a new energetic pathway from mussels to top predators. However, due to the short time elapsed after round goby establishment, we can only speculate on the species future impacts on Baltic food webs.     

Feeding preferences of an invasive Ponto‐Caspian goby for native and non‐native gammarid prey

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Residency Duration and Shelter Quality Influence Vibratory Signalling Displays in A Territorial Caterpillar

Residents are more likely to win territorial disputes than intruders. One explanation for this prior resident advantage is that residents place a higher value on the resource and are therefore more motivated to win. Although value asymmetry models of animal contests often assume that contestants use information about resource value, information on the proximate cues affecting territorial behaviour is often lacking. We use a simple model system – territorial behaviour in the masked birch caterpillar (Drepana arcuata) to identify factors that affect territorial behaviour. Late instar caterpillars occupy solitary silken leaf shelters, which they defend against wandering conspecifics with a vibratory display. We evaluated how a caterpillar identifies itself as the owner and the factors that influence a resident's motivation to signal. To do so, we conducted three experiments between size‐matched residents and intruders to assess how residency duration and shelter quality independently affected territorial displays during the early stages of a contest. Experiment 1 (Time Exp.) demonstrated that resident signalling rates increase with increased duration on the leaf prior to introducing the intruder. Residents also signal more than intruders after residency periods of 1–3 min and longer, demonstrating that residents gather information about resource value shortly after occupying a leaf. Experiment 2 (Squatter Exp.) aimed to disentangle the effects of time on the leaf and silk accumulation. Squatters (individuals in a shelter made by another) placed for 1–3 min on a leaf containing a full silk shelter signalled more to intruders than did caterpillars placed on a fresh leaf for 1–3 min. Experiment 3 (Shelter Removal Exp.) showed that residents whose shelters had been removed signal less than those occupying an intact shelter, despite an equal length of time investing in them. Our experiment is the first to covary both prior residency duration and territory quality, and we find that the motivation of caterpillars to signal is a function of both of these attributes.     

The Round Goby, Neogobius melanostomus, a Fish Invader on both sides of the Atlantic Ocean

During the past decade, a bottom-dwelling, aggressive, multiple-spawning fish, the round goby (Gobiidae: Neogobius melanostomus), has spread from its native region in the Ponto-Caspian throughout Europe and to the Laurentian Great Lakes in North America. An international workshop, held at the Hel Marine Station, Poland, was organized to summarize population features of the round goby. Common fish predators of round gobies in the Great Lakes and in native regions are obligate and facultative benthic fishes and occasionally, pelagic fishes. In contrast, the main predator of the round goby in the Gulf of Gdansk is the Great Cormorant (Phalacrocorax carbo). In the Great Lakes, round gobies have lead to the decline of mottled sculpin (Cottus bairdi) and logperch (Percina caprodes) and reduced the hatching success of native fishes by feeding on their eggs. In the Gulf of Gdansk, round gobies have increased in abundance, while three-spined sticklebacks (Gasterosteus aculeatus) have declined. Round gobies have a broad diet throughout their range; larger specimens are molluscivores. There are fewer species of parasites and lower infection rates of round gobies in recently colonized areas than in native areas. Overall, newly colonized round gobies in brackish waters and lakes are smaller, mature earlier, have a male biased operational sex ratio and are more short-lived compared with round gobies from marine (native) habitats.     

Factors affecting the outcome of territorial contests between hatchery and naturally reared coho salmon parr in the laboratory

In aquarium experiments using coho salmon as a model species, prior residents dominated intruders of the same size but intruders with a 6% length advantage were equally matched against prior residents. Prior winning experience (distinct from individual recognition) also strongly influenced competitive success and overcame a prior residence effect. Coho salmon reared in a hatchery dominated size-matched fish from the same parental population reared in a stream. Hatchery-reared salmon also dominated naturally spawned salmon, even when the wild salmon were prior residents. Thus the combined effects of greater size and rearing experience of hatchery-produced salmon were sufficient to overcome a wild salmon's advantage of prior residence. Efforts to rehabilitate salmonid populations must consider such behavioural interactions if displacement of wild fish is to be prevented.     

Some evidence for different ecological pressures that constrain male and female body size

The dwarf morph of the Lake Tanganyika cichlid Telmatochromis temporalis uses empty snail shells as shelters and breeding sites in shell beds, in which many empty shells exist. Here, we assessed selection forces regulating body size in this fish. Field observations showed that large males tended to have a greater number of females in their territories, suggesting that sexual selection favours large males. Nonetheless, a transplant experiment suggested that male body size was limited by the ability to hide in empty shells from large piscivorous fish. In females, the number of ovarian eggs increased with body size, suggesting that fecundity selection favours large females. However, females are smaller than males. Females spawned eggs close to the apices inside the shells. The small space there would decrease the risk of egg predation by egg predators, and small body size of females may be a result of adaptation to spawn eggs in the small, safe spaces. This study provides support for the idea that male and female body sizes have been limited by different ecological pressures (predation on adult fish in males, predation on eggs in females), which has not been reported previously in any animal.     

Predicting habitat use and trophic interactions of Eurasian ruffe,round gobies,and zebra mussels in nearshore areas of the Great Lakes

The Laurentian Great Lakes have been subject to numerous introductions of nonindigenous species, including two recent benthic fish invaders, Eurasian ruffe (Gymnocephalus cernuus) and round gobies (Neogobius melanostomus), as well as the benthic bivalve, zebra mussel (Dreissena polymorpha). These three exotic species, or “exotic triad,” may impact nearshore benthic communities due to their locally high abundances and expanding distributions. Laboratory experiments were conducted to determine (1) whether ruffe and gobies may compete for habitat and invertebrate food in benthic environments, and (2) if zebra mussels can alter those competitive relationships by serving as an alternate food source for gobies. In laboratory mesocosms, both gobies and ruffe preferred cobble and macrophyte areas to open sand either when alone or in sympatry. In a 9-week goby–ruffe competition experiment simulating an invasion scenario with a limited food base, gobies grew faster than did ruffe, suggesting that gobies may be competitively superior at low resource levels. When zebra mussels were added in a short-term experiment, the presence or absence of mussels did not affect goby or ruffe growth, as few zebra mussels were consumed. This finding, along with other laboratory evidence, suggests that gobies may prefer soft-bodied invertebrate prey over zebra mussels. Studies of interactions among the “exotic triad”, combined with continued surveillance, may help Great Lakes fisheries managers to predict future population sizes and distributions of these invasive fish, evaluate their impacts on native food webs, and direct possible control measures to appropriate species.     

Territorial behavior of the Korean wood-feeding cockroach, <Emphasis Type="Italic">Cryptocercus kyebangensis</Emphasis>

We investigated the territorial behavior of Cryptocercus kyebangensis from the viewpoint of offspring protection. Aggressive behavior against adult intruders occurred more frequently in resident adults with their young than those without their young. Resident adults without offspring more frequently attacked intruders of the same sex than those of the opposite sex, whereas those with offspring attacked regardless of the sex of adult intruders. In paired residents, cooperative territorial behavior occurred in a higher proportion in pairs with offspring than those without offspring. Responses of residents to nymphal intruders depended on the head capsule size of the intruders. Resident adults rarely attacked intruders smaller than their own offspring but frequently attacked larger intruders. Resident nymphs frequently attacked intruders smaller than themselves, whereas they escaped or showed jerking behavior when they encountered larger intruders. Electronic Publication     

Prior residence, body size and the dynamics of territorial disputes between male freshwater angelfish

Territorial interactions between pairs of size mismatched, sexually mature male angelfish Pterophyllum scalare were investigated in three different conditions: with the larger fish resident (the large resident condition), with the smaller fish resident (the small resident condition) and in a neutral territory (the neutral condition). In the two resident conditions, approximately half of the intruders had previously held territories and half had not. In all categories of fight, one fish showed submissive postures and lost the fight; eventual losers performed both attack and threat at a lower rate than eventual winners. Attack rate declined as the encounter progressed, while rate of performance of threat postures increased. In fights on neutral territories, the larger fish won all fights. In all fights with a resident-intruder asymmetry, the resident fish won the encounter, regardless of relative size. In eventual winners but not in eventual losers, levels of attack were lowest in the neutral encounters. In the small resident condition, levels of attack (corrected for activity of the resident) were lower in intruders that had previously held a breeding territory. Relative size influenced behaviour shown during fights, in that overall intensity was correlated negatively with size differential in all conditions. Thus although prior residence is the primary determinant of the outcome of territorial encounters in this species, both relative body size and prior possession of territory also influence the nature of the interaction.     

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.

     

Multiple cleaner species provide simultaneous services to coral reef fish clients

Cleaning symbioses on tropical coral reefs are typically documented between two species: a single client fish and one or more conspecific cleaners. However, multiple cleaner species living sympatrically in the Caribbean have been anecdotally reported to simultaneously clean the same client. Nothing is known about the patterns and processes driving these interactions, which may differ from those involving a single cleaner species. Here, we used remote underwater videography on three reefs in Honduras to record simultaneous cleaning interactions involving Pederson''s cleaner shrimp (Ancylomenes pedersoni) and cleaner gobies (Elacatinus spp.). A pilot study on adjacent shrimp and goby stations found interactions were always initiated by shrimp. A larger, multi-year dataset shows cleaner gobies joined 28% of all interactions initiated at A. pedersoni cleaning stations with cleaner gobies residing nearby. Client body size significantly predicted simultaneous cleaning interactions, with 45% of interactions simultaneous for clients greater than 20 cm total body length compared with only 8% for clients less than 20 cm. We also found that simultaneous cleaning interactions lasted over twice as long as shrimp-only interactions. We propose these novel multi-species interactions to be an ideal model system to explore broader questions about coexistence, niche overlap and functional redundancy among sympatric cleaner species.