Spawning season and migration of the myctophid fish <Emphasis Type="Italic">Diaphus theta </Emphasis>in the western North Pacific

 The spawning season, spawning grounds, and migration of the myctophid fish Diaphus theta were studied in the western North Pacific, based on seasonal sampling and estimation of hatching dates. The peak abundance of larvae was observed in July in the transition waters between the Oyashio and Kuroshio fronts. The spawning season ranged from late March to early September, with a peak from May to July. Larvae and juveniles <40 mm in standard length were distributed in the transition waters, whereas larger individuals were collected in the Oyashio and the Western Subarctic waters. These results indicate that this species undergoes a horizontal spawning migration from the Oyashio and Western Subarctic waters into the transition waters crossing the Oyashio front. Received: July 11, 2002 / Revised: October 2, 2002 / Accepted: October 15, 2002 Acknowledgments We are grateful to the captains, officers, and crew of FRV Hokko-Maru, FRV Tankai-Maru, and RV Hakuho-Maru for their assistance at sea. We thank Dr. H. Saito and Mr. H. Kasai for their cooperation in field sampling, and Dr. C. Sassa for his assistance with larval fish taxonomy. Correspondence to:Masatoshi Moku     

Combining limnological and palaeolimnological approaches in assessing degradation of Lake Pskov

Historical spawning grounds for sea lampreys (Petromyzon marinus L.) in most Portuguese river basins are becoming inaccessible due to the construction of impassable dams and/or weirs. Studies like the one described in this paper are particularly important in areas like the Vouga river basin, where there is a considerable fishing effort from both professional fishermen and poachers. In fact, for management and conservation purposes, it is important to clarify several aspects of the sea lamprey spawning run in this particular watershed. Therefore, a total of 30 radio tagged, migrating sea lampreys were released in the River Vouga and some of its main tributaries during 2004 and 2005. Results from the tracking sessions were used to determine the effect of poaching on the spawners’ population and the characteristics of the resting sites used during the upstream movement. The rivers’ stretches were also characterized according to the type of substrate present in the riverbed and flow type, in order to determine its aptitude to constitute appropriate spawning habitats for sea lampreys. We have identified in the upstream stretches of River Vouga, and in the tributary River Caima, characteristics that are particularly suitable for the construction of nests by the spawners. Presence of larvae provided evidence that spawners migrated into the River Vouga’s upper reaches. However, abundance and age class diversity appeared to be higher downstream of Sernada and Carvoeiro weirs, corroborating the telemetry data which suggested difficulty in passing these obstacles during low precipitation years. In the River Caima, migration was most predictable, which might be related to daily water releases from a small hydropower dam at dusk, that stimulated the lampreys to resume migration. Poaching has had a considerable negative effect on the success of the tagged lampreys’ migration: 76% of the animals released during 2005 were captured. Electronic supplementary material  Supplementary material is available for this article at and accessible for authorized user     

Sperm allocation pattern during a reproductive season in the copulating marine cottoid species, <Emphasis Type="Italic">Alcichthys alcicornis</Emphasis>

The sperm allocation pattern of a copulating marine cottid fish, Alcichthys alcicornis, was investigated. A total of 86 mating events using six males were conducted in aquarium tanks over 10 days, and in 36 of them, spermatozoa were collected using a false copulation method. Males released 3–8 × 10⁸ spermatozoa in early events, with the number decreasing gradually during subsequent mating events. This sperm allocation was represented as an “early investment and tapering” pattern. It was discussed why males have significantly higher sperm release in early spawning events. The reproductive behavior consists of spawning and subsequent copulation. Spermatozoa have the ability to fertilize eggs from multiple clutches, and in earlier produced clutches the level of sperm competition should be relatively low. In addition, if early spawn happens to be the first spawn with a female, spermatozoa that are released into the water column after spawning are responsible for fertilizing the female’s first clutch. The probability of this occurring should decrease dramatically as the season progresses, due to the highly synchronous seasonal spawning of females. All of these factors should select for high sperm numbers in early ejaculates. Based on such reproductive ecology of A. alcicornis we hypothesize that this sperm allocation pattern is an adaptive reproductive strategy in response to egg availability and sperm competition occurring within the ovarian cavity.     

A new direct method of stock assessment of the loliginid squid

Hydroacoustic research conducted on chokka squid (Loligo reynaudi d’Orbigny, 1845), off the east coast of South Africa from 1994–2005, has led to the development of an innovative stock assessment technique, perhaps applicable to all loliginids that migrate inshore to spawn. This technique combines hydroacoustic biomass estimates made on the spawning concentrations inshore, and minimum biomass estimates made both inshore and offshore using demersal surveys employing the swept-area method. The hydroacoustic estimate uses an improved method to obtain target strength measurements, and squid concentrations are individually mapped from a small boat with a towed transducer. This method may be used even during intense fishing operations because of the manoeuvrability of the small boat inside a tight cluster of fishing vessels. Biomasses of the individual concentrations are then summed. The inshore biomass, also includes dispersed, mature squid migrating between concentrations, this is assessed using a concentration stability factor. The biomass of dispersed squid offshore is again calculated using the swept-area method, a well known demersal survey methodology. The biomass of concentrated (spawning) squid offshore is calculated using the same proportions between concentrated and dispersed squid which were found inshore. All four components are then summed to calculate the total biomass. The result obtained is subject to the effect of complex temporal dynamics, as new animals are recruited to the adult pool and those recently assessed migrate to other sectors of the distribution area.     

Quantitative composition,distribution, and feeding of large jellyfish (<Emphasis Type="Italic">Scyphozoa et Hydrozoa</Emphasis>) on the West Kamchatka shelf in summer

The total biomass of jellyfish on the shelf of the eastern Sea of Okhotsk in the summer is estimated as 1672700 tons according to the results of hydroacoustic measurements and 901000 tons by the method of squares. The use of hydroacoustic technologies makes evaluation of the actual stock and range of medusae more accurate, and the further enhancement and perfection of the hydroacoustic method based on multifrequency measurements enables one to obtain more reliable estimates. A significant increase (nearly 25 times) of the total jellyfish biomass takes place in the summer period. Cyanea prevailed in biomass in the spring and Chrysaora melanaster prevailed in the summer. Some species showed considerable expressed spatial differentiation of distribution and affinity to certain environmental conditions. The studied species were almost exclusively zoophages. Their algal diet consisted mainly of diatom algae. Scyphomedusa’s diet mainly included the so-called “peaceful” zooplankton, viz., euphausiids and copepods (as a rule, over 50% the mass), at the same time carnivorous zooplankton, saggits, amphipods, and small medusae also formed a substantial share of their diet. One individual of the predominant jellyfish species consumes a total of 6.1 to 70.5 kcal during its lifecycle, which corresponds to 79.1–513.0 g of raw organic material, assuming 70% assimilability. The relatively low demand for food of this sort can be explained by the low caloric value of the jellyfish body, 96–97% of which consists of water. The distribution and composition of the jellyfish prey show that scyphomedusae exert the greatest influence on the nekton community, as they concentrate in the shelf area of the eastern part of the sea, at walleye pollock spawning sites. There the larvae of bottom invertebrates, including commercially valuable organisms, such as crab and shrimp, are also consumed. In the summer, jellyfish eat nearly 100 billion eggs and 20 billion larvae of walleye pollock, as well as 130 billion decapod (mostly crab) larvae each day, which corresponds to 0.03% of the eggs and 0.003% of the larvae of walleye pollock and 0.003% of the decapod larvae in the estimated stock.     

Patterns of migration between feeding and spawning sites in a coral reef surgeonfish

Many coral reef fishes exhibit regular localised migrations between feeding and spawning areas, but the factors affecting these migration patterns, such as the distance, frequency and spawning site fidelity are poorly understood. The aim of this study was to investigate the patterns of migration to spawning sites of the surgeonfish, Ctenochaetus striatus (Acanthuridae). We explored relationships amongst an individual’s size and sex, the distance and frequency it migrated from its feeding area to spawning sites, fidelity to particular spawning sites and the number of individuals that aggregated to spawn. In order to achieve this, 406 C. striatus were captured and tagged on inshore reefs in Kimbe Bay (5°30′S 150°6′E), New Britain, Papua New Guinea. Tagged individuals were consistently observed within spatially discrete but overlapping feeding areas (maximum diameter averaging <13 m). The mean distance migrated was 58 m (ranging from 2 to 291 m). No tagged individuals were witnessed spawning at more than one site. Whilst most individuals (n = 88) migrated to the spawning site that was closest to their feeding areas, those that migrated to sites further away (n = 9) always spawned at sites where the number of conspecifics aggregating was larger. Neither the size nor the sex of individuals limited migration distance. However, males migrated significantly more frequently than females (on average once every 2 days vs. once every 3 days), and migration frequency was positively correlated with size in females. Migration distance did not affect the frequency with which individuals spawned. Whether patterns of migration are determined by cost-benefit optimisation, tradition, or an alternative mechanism is unknown.     

Quantitative Links Between Pacific Salmon and Stream Periphyton

Species’ impacts on primary production can have strong ecological consequences. In freshwater ecosystems, Pacific salmon (Oncorhynchus spp.) may influence stream periphyton through substrate disturbance during spawning and nutrient subsidies from senescent adults. The shape of relationships between the abundance of spawning salmon and stream periphyton, as well as interactions with environmental variables, are incompletely understood and may differ across the geographic range of salmon. We examined these relationships across 24 sockeye salmon (Oncorhynchus nerka) spawning streams in north-central British Columbia, Canada. The influence of salmon abundance and environmental variables (temperature, light, dissolved nutrients, water velocity, watershed size, and invertebrate grazer abundance) on post-spawning periphyton abundance and nitrogen stable isotope signatures, which can indicate the uptake of salmon nitrogen, was evaluated using linear regression models and Akaike Information Criterion. Periphyton nitrogen stable isotope signatures were best described by a positive log-linear relationship with an upstream salmon abundance metric that includes salmon from earlier years. This suggests the presence of a nutrient legacy. In contrast, periphyton abundance was negatively related to the spawning-year salmon density, which likely results from substrate disturbance during spawning, and positively related to dissolved soluble reactive phosphorus prior to spawning, which may indicate phosphorus limitation in the streams. These results suggest that enrichment from salmon nutrients does not always translate into elevated periphyton abundance. This underscores the need to directly assess the outcome of salmon impacts on streams rather than extrapolating from stable isotope evidence for the incorporation of salmon nutrients into food webs.     

Assessing the effect of Separation Levee Project on Chinese sturgeon (<Emphasis Type="Italic">Acipensor sinensis</Emphasis>) spawning habitat suitability in Yangtze River,China

Chinese sturgeon (Acipenser sinensis) is a protected anadromous fish species. The migration pattern of the fish has been blocked by the construction of Gezhouba Dam, reducing the natural spawning site length to less than 7 km along the Yangtze River. However, the fish has since established an alternative spawning ground in the narrow reach downstream of Gezhouba Dam. To enhance navigation, a Separation Levee Project (SLP) was implemented in the new-found spawning habitat of the fish. To therefore evaluate the effect of the SLP on Chinese sturgeon spawning habitat suitability, the conditions in the spawning habitat were simulated using River2D (a two-dimensional hydrodynamic model). Two main approaches (habitat kinetic energy and circulation metrics) were used in the simulation. The study showed that SLP only slightly changed the physical conditions in the spawning habitat. Using hydrodynamic simulation, the weighted usable area (WUA) before and after the SLP construction was also computed and habitat preference curve developed for water depth and velocity. On the average, SLP reduced WUA—a finding that was consistent with field-measured data. Based on WUA, the habitat conditions were more sensitive to SLP proximity than metrics based on velocity gradients. SLP posed detrimental impacts on the suitability of spawning habitats of Chinese sturgeon. The findings in this study provide further basis for the protection and restoration of Chinese sturgeon spawning habitats in especially the lower reach of Yangtze River.     

Resilience of an isopod population (<Emphasis Type="Italic">Cyathura carinata</Emphasis>) to multiple stress factors in a temperate estuarine system

The Mondego Estuary (Portugal) underwent major modifications in the late 1980s, because of eutrophication, for which it was developed a restoration programme that has been effective in recovering seagrass beds. Recently, this system has been affected by successive weather extremes: floods, droughts, and heat waves. The effect of these multiple stressors on a Cyathura carinata (Isopoda) population has been evaluated, through a long-term data series (16 years) in a seagrass bed, an intermediate area, and a sand flat. A well age-structured and abundant population of C. carinata was observed in the sand flat, in contrast with the seagrass bed and intermediate area. After introducing the restoration measures, C. carinata’s abundance increased even in the stations where it was formerly scarce. This recovery was compromised by successive extreme events. C. carinata declined severely in terms of total abundance and growth production during the 2003 summer heat wave, at the sand flat, also endangering its reproductive success. Floods and droughts affected dispersion of the individuals, which was reflected in the population’s density and production. Nevertheless, this isopod had great resilience, recovering successfully after disturbances. In contrast, the population of C. carinata had less resilience in the seagrass bed and intermediate area, being unable to recover after a succession of disturbances, which may threaten the success of this species in the system. Because the frequency and intensity of weather extremes are expected to continue to increase in a scenario of global climate change, their repercussions will affect such populations, which will have difficulty surviving. This will have severe implications on the functioning of the whole ecosystem.     

DELIMITING SPAWNING AREAS OF WEAKFISH CYNOSCION REGALIS (FAMILY SCIAENIDAE) IN PAMLICO SOUND,NORTH CAROLINA USING PASSIVE HYDROACOUSTIC SURVEYS

ABSTRACT

Exact locations of spawning areas used by marine fishes are needed to design marine reserves and estimate spawning stocks. The location of spawning areas of soniferous fishes such as weakfish Cynoscion regalis can be determined by means of passive hydroacoustic surveys. We conducted nocturnal hydrophone surveys at 12 locations in Pamlico Sound in May of 1996 and 1997. Digital audio tapes were made of weakfish “purring” sounds, the tapes were analyzed spectrographically and compared with ichthyoplankton surveys taken at the same stations and times. All weakfish “purring” sounds were recorded at stations near inlets. Maximum sound pressure levels recorded after sunset were 127 dB (re 1 (μPa) for individual weakfish, but reached a maximum of 147 dB when weakfish and other fish were producing sounds simultaneously. The maximum distance that an individual weakfish “purr” can be detected above the background sound, assuming a cylindrical spreading model, is approximately 50 m. There was a strong association (r = 0.78) between the log₁₀— transformed abundance of early-stage sciaenid-type eggs and maximum sound pressure levels, with the greatest numbers occurring at the inlet stations. These results suggest that passive hydroacoustic surveys can be used to delimit spawning areas for conservation and management purposes.     

Pollination effectiveness and pollinator importance in a population of Heterotheca subaxillaris (Asteraceae)

 Assessing the relative contributions to seed set for each of a plant species’ floral visitors provides an indication of the relative influence of these visitors on the plant’s reproductive success. This study examined pollinator activity and seed set in a population of Heterotheca subaxillaris, a species that exhibits a floret dimorphism (heads bearing disk and ray florets), and that is visited by both generalist foragers and specialist bees. Visits by nine bee genera and one genus of skipper were recorded in the study population. During the period of study, these insects varied in their relative abundance, in their foraging activity on a head, and in their pollination effectiveness. The pattern of pollination effectiveness shown by the different pollinators was similar for both floret types, although seed set was higher overall for ray florets. Pollinator importance, calculated as the product of pollination effectiveness and relative abundance, was dictated by a pollinator’s relative abundance. The single specialist bee species observed in the study population proved to have neither higher pollination effectiveness nor higher pollinator abundance than other pollinators. This would suggest that H. subaxillaris is not under strong selective pressure to co-specialize with its specialist visitor. Received: 21 December 1995 / Accepted: 20 June 1996     

Influence of modern irrigation,drainage system and water management on spawning migration of mud loach, <Emphasis Type="Italic">Misgurnus anguillicaudatus</Emphasis> C.

We investigated the influence of human activities on the spawning migration of the mud loach, Misgurnus anguillicaudatus. Mud loaches migrate from rivers to paddy fields for spawning. In this study, the number of adult loaches in paddy fields showed a peak in late May and rapidly decreased in July. In early July, water is discharged from paddy fields in order to allow the fields to dry for a few weeks, and water is again supplied later. The spawning season of the loaches probably spans from mid May to August; this was concluded by observing physiological indicators such as the gonad somatic index, oocyte maturity and serum levels of testosterone, estradiol-17β and 17α, 20β-dihydroxy-4-pregnen-3-one. When the loaches mature, they migrated to paddy fields if they could swim upward through the drainage ditches. However, the drainage ditches of modern paddy fields were reconstructed to deepen the water levels. The artificial drought of the rice culture management led to a discharge of water and loaches from the paddy fields during their spawning season. These results indicate that modern rice cultivation causes obstruction of the migration route and loss of spawning grounds; consequently, loaches use limited space and period of the paddy field for their spawning.     

A flyway perspective on food resource abundance in a long-distance migrant, the Eurasian teal (Anas crecca)

Two frequent assumptions about the evolution of long-distance migration in birds are that they travel long distances annually to reach food-rich areas for breeding, and that they time their migratory journey to be at staging sites when the latter provide the best feeding conditions. These assumptions have rarely been properly tested, and there is no study in which a species’ major food types have been measured by standardized methods throughout a flyway and over a large part of the year. We here present such data for Eurasian teal (Anas crecca), converted to a common energetic currency, and collected at wintering, spring staging and breeding sites. Teal did not time migration to maximize local food abundance; most birds left wintering and spring staging sites before a sharp increase in invertebrate food abundance occurred. On the other hand, hatching of ducklings coincided with a peak in invertebrate food abundance on boreal breeding lakes. Mean overall food abundance (invertebrates and seeds combined) did not differ between wintering sites in southern France and breeding sites in northern Sweden at the time of breeding. Our results are inconsistent with the hypothesis that long-distance migration in dabbling ducks has evolved because adult birds gain an immediate pay-off in increased food abundance by flying north in spring. However, our data confirm a selective advantage for breeding at higher latitudes, because hatching of ducklings may coincide with a peak in invertebrate emergence and because longer days may increase the duration of efficient foraging.     

Spatial and temporal patterns of movement and migration at spawning aggregations of red hind, <Emphasis Type="Italic">Epinephelus guttatus</Emphasis>, in the U.S. Virgin Islands

This study compared the spatial and temporal patterns of red hind, Epinephelus guttatus, movement and migration from annual spawning aggregations on St Thomas (STT) and St Croix (STX), United States Virgin Islands. Around STT E. guttatus migrated 6–33 km from a functional spawning migration area of 500 km² and around STX E. guttatus migrated 5–18 km from an area of 90 km². Similarities between sites were found in regards to timing of movement, temporal and spatial changes in sex ratios, annual and lunar predictability and were synchronized with environmental cues. E. guttatus spawning aggregations in the Virgin Islands occur between the winter solstice (i.e., after December 20) and about February 20 of any year and show a distinctive peak 20–40 days after the winter solstice. Spawning typically occurred during periods of declining seawater temperature and slacking currents within a temperature range of 26–27.5°C and current speed of 2.5–3.5 cm s⁻¹. Males arrived early to spawning sites and stayed longer than females. These gender-based behavioral patterns are important to E. guttatus reproductive dynamics and must be factored into future studies and the design of fisheries regulations to ensure sustainability of spawning aggregation sites. The predictability of E. guttatus spawning aggregations relative to the winter solstice will be extremely beneficial for defining the temporal and spatial aspects of area closures. The consistency and synchrony of movement and migration will improve both the efficiency of planning research and monitoring programs and directing enforcement activities during critical time periods. Applying this knowledge strategically will maximize the limited resources available for research and enforcement and lead to greater protection of spawning aggregations.     

Effect of spatial differences in growth on distribution of seasonally co‐occurring herring Clupea harengus stocks

The mechanisms most likely to determine the distribution of the two major herring Clupea harengus stocks in their common early summer feeding ground in the eastern North Sea, Skagerrak and Kattegat were investigated through analysis of acoustic survey data from six consecutive years. No change was detected in biomass of North Sea autumn spawning C. harengus (NSAS) over time, whereas the biomass of western Baltic spring spawning C. harengus (WBSS) declined severely. Analyses of centre of abundance by stock showed no change in NSAS distribution, whereas the WBSS changed to a more western distribution over time. Contrary to previous perception of the juvenile migration, NSAS were found to leave the study area at the age between 1 and 2 years and WBSS 1 year olds were encountered in the Skagerrak. The estimated parameters of von Bertalanffy growth equations showed marked differences between areas with fish in the eastern part of the area having the lowest size at age at all ages. Further, their growth conditions appeared to deteriorate progressively over the period studied. Both NSAS and WBSS showed the highest condition in the North Sea and Skagerrak while condition was substantially lower in age Kattegat. The westward movement of spring spawners over time suggests that growth rate and possibly density of conspecifics influence the migration pattern and distribution of C. harengus in the area. In contrast, there was no evidence to suggest that distribution was constant over time within stocks or that distribution reflected size‐dependent limitations on migration distance.     

Factors affecting large-scale distribution of the Bonelli’s eagle <Emphasis Type="Italic">Aquila fasciata</Emphasis> in Spain

We analyzed the environmental determinants of the regional distribution of Bonelli’s eagle (Aquila fasciata) in Spain (ca. 500,000 km²), taking into account its frequency of occurrence on UTM blocks of 50 × 50 km. We found that the distribution pattern of Bonelli’s eagle was a highly predictable phenomenon based on climate, vegetation and interspecific relationships. The proportion of sunny, anticyclonic days, per year (i.e., high levels of solar radiation) was the main environmental predictor explaining the distribution and abundance pattern of the Bonelli’s eagle. Sparse plant formations (mainly shrublands) had also a positive effect, while altitude, agricultural land and deciduous forests had a negative influence. The relative abundance of one of its main preys, the Red Partridge (Alectoris rufa), favored the probability of occurrence of Bonelli’s eagle, but only in the less sunny areas located in the north of Spain. The relative abundance of its main competitor, the Golden eagle (Aquila chrysaetos), was slightly but positively correlated with both the distribution and the abundance of the Bonelli’s eagle in the Spanish portion of the Iberian Peninsula, probably due to similarities in their habitat preferences. Finally, we did not find further regional effects of the variables describing the degree of human pressure (density of roads, urban cover, and length of power lines).     

Investigations of spawning migration and assessment of abundance of the Kamchatka steelhead (<Emphasis Type="Italic">Parasalmo mykiss</Emphasis>) from the Utkholok River by means of Didson dual-frequency identification sonar

Quantative estimates of abundance of anadromous Kamchatka steelhead Parasalmo mykiss in the period of its maximum spawning migration in the shallow Utkholok River by the instrumental method using Didson dual-frequency identification sonar are presented. Equipment and methods are described for the stationary observation of spawning run of Kamchatka steelhead through the river cross section and mobile surveys of the water area of the Utkholok R. As a result of stationary observations, the abundance and size composition of Kamchatka steelhead migrating to the upper reaches of the river are determined. Peculiarities of the diurnal dynamics of migration of Kamchatka steelhead characterized by two sharp peaks of run intensity in the morning and evening were revealed. During the period of observations from September 22 to October 20, 2007, in the seasonal dynamics of migration two peaks of intensity of run of Kamchatka steelhead depending on an abrupt decrease in water temperature by 2°C were also registered. Mobile surveys supplied data on the abundance and size composition of Kamchatka steelhead in the surveyed water area of the river, on its spatial distribution, and dynamics of accumulation of fish in the river. The abundance of Kamchatka steelhead in the river almost doubled every three-four days and the spatial distribution was aggregated, producing maximum concentration mainly in areas of increasing depth and channel meandering before and after riffles. Comparison of the results of assessment of abundance of Kamchatka steelhead that arrived for spawning in 2006 and 2007 showed significant fluctuations in population abundance.     

Lake trout (<Emphasis Type="Italic">Salvelinus namaycush</Emphasis>) spawning behaviour: the evolution of a new female strategy

A video camera inside an underwater housing was used to record the spawning activities of lake trout, Salvelinus namaycush, in Kushog Lake, Ontario (Canada). Contrary to the commonly accepted belief describing lake trout as the only salmonine to spawn exclusively at night, the recordings were taken during the day. Lake trout spawning behaviour is described in detail and compared with other salmonine species. The loss of female “nest-building” and assumption of a novel itinerant strategy during which females travel across the breeding grounds accompanied by a group of males, is considered to be an important change in the evolution of the lake trout’s unique mode of spawning.     

The significance of sound interception to males of the bicolor damselfish,Pomacentrus partitus,during courtship

Synopsis Playback experiments conducted in the field and the laboratory demonstrated the use of sound interception in bicolor damselfish,Pomacentrus partitus. Two courtship sounds produced by male bicolors: the ‘chirp’ which occurs at the initiation of courting and the ‘grunt’ which occurs near the termination, just prior to spawning, were found to hold different meanings to intercepting male competitors. Males responded to grunt playback with directional swimming towards the sound source and increased courtship behavior. No directional response was observed during chirp playback. The grunt sound appears to indicate that a spawn-ready female is present near the sound source. Males therefore move towards it, likely to interfere with the imminent spawning or to gain a possible spawning partner. Such is not observed in response to chirps, as little advantage would be gained by moving to the source of a sound which is known to serve as an ‘advertisement’ or ‘territorial keep-out’ signal. Thus, upon intercepting a neighbor's chirps, males exhibit courting within their own territory, or directly court a nearby female.