A major increase in snake pipefish (<Emphasis Type="Italic">Entelurus aequoreus</Emphasis>) in northern European seas since 2003: potential implications for seabird breeding success

Since the early 2000s routine fish surveys have recorded increasing numbers of snake pipefish, Entelurus aequoreus, in the northeast Atlantic. Fishermen and divers have also commented on this increase and pipefish have started to appear in the diet of seabirds and other marine predators. This paper collates information from these diverse sources and assesses the current status of snake pipefish. We found compelling evidence of a dramatic increase in the abundance of snake pipefish starting around 2003 and continuing up to the present (2006) and a range expansion northwards to Spitzbergen and the Barents Sea. Since 2004 snake pipefish have been increasingly recorded in the diet of many species of seabird breeding in colonies around the coast of the UK, and in Norway, Iceland and the Faeroe Islands. Information on the nutrient value of snake pipefish is currently lacking but their rigid, bony structure makes them difficult for young seabirds to swallow and there are numerous records of chicks choking to death. Thus, in the case of avian predators during the breeding season, it appears unlikely that increased abundance of snake pipefish will provide a useful alternative prey. The reason for the rapid and dramatic increase in numbers of snake pipefish is currently unclear but such events are characteristic of marine ecosystems and will almost certainly have an effect on food web dynamics.     

Contrasting foraging tactics by northern gannets (<Emphasis Type="Italic">Sula bassana</Emphasis>) breeding in different oceanographic domains with different prey fields

In order to forage and to provision offspring effectively, seabirds negotiate a complex of behavioural, energetic, environmental and social constraints. In first tests of GPS loggers with seabirds in North America, we investigated the foraging tactics of free-ranging northern gannets (Sula bassana) at a large and a medium-sized colony that differed in oceanography, coastal position and prey fields. Gannets at Low Arctic colony (Funk Island) 50 km off the northeast coast of Newfoundland, Canada provisioned chicks almost entirely with small forage fish (capelin Mallotus villosus, 89%), while at boreal colony (Bonaventure Island) 3 km from shore in the Gulf of St. Lawrence, Quebec, Canada, large pelagic fish dominated parental prey loads (Atlantic mackerel Scomber scombrus 50%, Atlantic herring Clupea harengus 33%). Mean foraging range and the total distance travelled per foraging trip were significantly greater at the larger inshore colony (Bonaventure) than at the smaller offshore colony (Funk Island; 138 and 452 km vs. 64 and 196 km, respectively). Gannets from Funk Island consistently travelled inshore to forage on reproductive capelin shoals near the coast, whereas foraging flights of birds from Bonaventure were much more variable in direction and destination. Birds from the Low Arctic colony foraged in colder sea surface water than did birds from the boreal colony, and dive characteristics differed between colonies, which is concordent with the difference in prey base. Differences between the colonies reflect oceanographic and colony-size influences on prey fields that shape individual foraging tactics and in turn generate higher level colony-specific foraging “strategies”.     

Primary productivity and related data from tropical and subtropical marine sediments

Measurements in situ were made of oxygen exchange and ¹⁴C fixation in calcareous sediments off the coast of Florida, USA, and at various sites in the Caribbean Sea at depths to 60 m. Sediment samples were analyzed for total organic carbon, nitrogen and photosynthetic pigments. In some cases, measurements were made of the pH of sediment interstitial water and its concentration of total CO₂. In the Caribbean Sea, routine hydrological sampiing was undertaken and in situ plankton productivity determined.Contribution No. 1522 from the University of Miami, Rosenstiel School of Marine and Atmospheric Science, 10 Rickenbacker Causeway, Miami, Florida 33149, USA.     

Microsetella norvegica: a rare report of a potentially abundant copepod on the Scotian Shelf

Fine mesh (0.080 mm) zooplankton samples were collected along an onshore-offshore transect during three cruises (February to May, 1983) off southwest Nova Scotia, Canada. The small harpacticoid copepod Microsetella norvegica was numerically the dominant copepod during the winter (February) cruise, where it composed up to 50% of the total zooplankton, and was consistently among the three most abundant copepods during the spring cruises. Due to its small size (<0.45 mm in length), M. norvegica has rarely been reported from plankton surveys of the northwest Atlantic. However, existing reports indicated it varies interannually from being a rare to an abundant member of the zooplankton community in the northwest Atlantic.     

Effect of low dissolved oxygen concentrations on behavior and predation rates on red sea bream Pagrus major larvae by the jellyfish Aurelia aurita and by juvenile Spanish mackerel Scomberomorus niphonius

A shift in outcomes of predator-prey interactions in plankton community may occur at sublethal dissolved oxygen concentrations that commonly occur in coastal waters. Laboratory experiments were conducted to investigate how a decline in dissolved oxygen concentration alters the predation rate on fish larvae by two estuarine predators. Behavior and consumption of larval fish by moon jellyfish Aurelia aurita (103.1±12.4 mm in bell diameter) and by a juvenile piscivore, Spanish mackerel Scomberomorus niphonius (30.1±2.1 mm in standard length: SL), were observed under four oxygen concentration treatments (1, 2 and 4 mg l^–1 and air-saturated: 5.8 mg l^–1). Larvae of a coastal marine fish species, red sea bream Pagrus major (7.21±0.52 mm SL), were used as prey for the experiment. Bell contraction rate of the jellyfish did not vary among the oxygen concentrations tested, indicating a tolerance to low oxygen concentration. Gill ventilation rate of the Spanish mackerel increased and swimming speed decreased as the oxygen concentration decreased, indicating that oxygen concentrations 4 mg l^–1 are physiologically stressful for this species. The number of larvae consumed in 15 min. by jellyfish increased whereas those consumed by Spanish mackerel decreased with the decrease in oxygen concentration. Low oxygen concentrations that are commonly observed in coastal waters of Japan during summer have the potential to increase the relative importance of jellyfish as predator of fish larvae and to change the importance of alternative trophic pathways in estuarine ecosystems.Communicated by T. Ikeda, Hakodate     

Polychlorinated-dibenzo-p-dioxins, -dibenzofurans and -dioxin-like polychlorinated biphenyls in aquatic organisms from lake Kasumigaura,Japan

Polychlorinated-dibenzo-p-dioxins (PCDDs), -dibenzofurans (PCDFs) and -dioxin-like polychlorinated biphenyls (DLPCBs) were determined in aquatic wildlife of Kasumigaura Lake (KUL), Japan from 1978 to 2001. Two plankton species elucidated several fold greater concentrations (2400–7800) than small tiger fish (310–6500), shrimp (160–1100), and three large fish namely, black bass (120–240), carp (94–120) and mullet (54) on pg/g fat. PCDD homologues were predominant accumulants with >68–<95% contribution and remaining was shared by PCDFs. Accumulation, non-ortho DLPCBs in plankton (3500–9200), shrimp (1600–8600) and small fish (2000–8800) and large fish (1300–3900) on ng/g fat basis were several orders magnitude greater than PCDD/DFs. The mono-ortho DLPCBs in large fish species were 23?000–83?000?ng/g fat and which it is accounted 94.3–95.9% the total dioxin-like PCBs accumulation. Temporal trends of PCDDs, PCDFs, non-ortho PCBs and TEQ in shrimp and small tiger fish were varied. The mullet had minimum toxic equivalency “TEQ” (14?pg?TEQ/g) followed by plankton Neomysis intermedia (25?pg?TEQ/g), small tiger fish (mean: 33, ranges 14–66?pg?TEQ/g), plankton Cyclopus vicinus (34?pg?TEQ/g), carp (mean: 35, ranges 32–38?pg?TEQ/g), shrimp (mean: 38, ranges 11–68?pg?TEQ/g) and black bass (mean: 59, ranges 38–79?pg?TEQ/g) on fat basis. In all the samples, PCDD was predominant TEQ contributor followed by PCDFs and DLPCBs. The contribution of mono-ortho DLPCBs to the total TEQ was 0.52–0.92 in large fish.     

Ecology of a Caribbean coral reef. The Porites reef-flat biotope: Part II. Plankton community with evidence for depletion

A quantitative assessment of drifting net plankton crossing a reef-flat biotope was obtained on a Caribbean coral reef. The spatial distribution and abundance of plankton were sampled to provide estimates of the removal of this potential food resource by suspension-feeding populations. Sampling was largely confined to the reef flat and adjacent waters of Laurel Cay, a flourishing coral reef present on the insular shelf off southwestern Puerto Rico. A prior study provides information on the meteorological and hydrographic characteristics of this area. Evidence for plankton accrual was found in the quantitative depletion of qualitatively similar populations sampled downstream of densely populated reef communities. Numerically, the diatom crop was reduced by 91% and zooplankton by 60% in water streaming off the reef. Significant diel and seasonal variations in plankton abundance were obcerved, as well as notable differences in volume flow, the latter closely related to the local wind regime. A time course of net plankton accrual was calculated, taking into account these various factors. During the summer season (July–August), when zooplankton was relatively abundant and water movement over the reef vigorous, the total gain from plankton reached 0.25 gC/m²/day; 75% of this occurred during a 4 h period at sunrise and sunset. Plankton retained on the reef flat in January of February and in September was around 0.1 g C/m²/day. Zooplankton biomass contributed the greatest share, exceeding that of diatoms by a factor of 10 during the day and 42 in the early evening. A mean annual accrual of 0.18 g C/m²/day is equivalent to 4 to 13% of net community metabolism.     

The effect of perceived female parasite load on post-copulatory male choice in a sex-role-reversed pipefish

The last several decades of research in behavioral ecology have resulted in a deeper appreciation of post-mating processes and sexual conflict in sexual selection. One of the most controversial aspects of sexual selection is cryptic mate choice. Here, we take advantage of male pregnancy in a sex-role-reversed pipefish (Syngnathus typhle) to quantify cryptic choice based on perceived parasite load and other sources of variance in female fitness. Studies have shown that S. typhle males preferentially mate with females with lower parasite loads and that a male’s perception of female parasite load can be altered by tattooing females. We manipulated the apparent parasite load of females in controlled mating experiments to test the hypothesis that post-copulatory sexual selection is dependent on a male’s perception of female parasite load in pipefish. Our results provided no evidence for cryptic male choice based on perceived female parasite load. However, we found evidence that eggs from larger females were more likely to result in viable offspring than eggs from smaller females and that the first female to mate with a male transferred more eggs per copulation on average. Overall, our results show that potential for post-copulatory sexual selection does exist in pipefish, but the male’s perception of female parasite load does not play a major role in this process.     

Ecological investigations of the late-stage phyllosoma and puerulus larvae of the western rock lobster Panulirus longipes cygnus

The distribution and abundance of the late-stage phyllosoma larvae of Panulirus longipes cygnus George and the distribution and densities of the final larval stage, the puerulus, both in the plankton and at settlement along the coast, were investigated. A total of 3,617 late-stage phyllosoma (Stages VI to IX) and 301 puerulus larvae were caught at 187 plankton stations during the July to November periods 1974, 1975 and 1976 off the west coast of Australia between 29°00 to 32°30S and 113°30 to 115°00E. The depth range sampled was 0 to 35 m on the continental shelf and 0 to 90 m off the shelf. During onshore/offshore cruises with similar sampling effort on and off the shelf, 1,169 late-stage phyllosoma larvae were taken, of which only 9 were caught on the shelf, and these near the outer edge. A series of cruises sampling two areas beyond the shelf near 29°30 and 32°00S yielded 2448 late-stage phyllosoma, with greater densities of larvae in the northern location. The settlement of puerulus-stage larvae along the coast in the same geographical range was also greater in the north than in the south. The data from the onshore/offshore cruises showed a definite effect of moon phase on numbers of puerulus larvae caught on the shelf, with higher catches near new moon. The low numbers of puerulus larvae (usually 0, 1 or 2 individuals) caught at all stations showed that the puerulus stage is sparsely distributed in the plankton. Fewer puerulus larvae were present at the surface than at lower depths, but it was not possible to determine a depth preference for the puerulus between 10 m and the lowest depths sampled because of the low catch numbers. No relationships were found between puerulus larvae density and surface-water temperature, salinity, or plankton biomass at each station. Data on the larval distributions indicate that, near the end of their planktonic existence, the majority of the late-stage phyllosoma larvae of P. longipes cygnus are not carried onto the shelf, where mixing of oceanic and continental shelf waters occurs only on the outer third, but are transported southward by oceanic circulation beyond the shelf. The puerulus moults from the last phyllosoma stage beyond the shelf and completes the larval cycle by swimming across the shelf and settling in the shallow reef areas.     

Biomass and production in polar planktonic and sea ice microbial communities: a comparative study

Algal and bacterial biomass and production were measured in the plankton, platelet ice and congelation ice communities at one station in McMurdo Sound, Antarctica during September and October 1986. Bacterial abundances and particulate organic carbon and nitrogen were 10 to 100 times greater in the plankton than in the sea ice, whereas the chlorophyll a concentrations in the plankton and sea ice microbial communities (SIMCO) were similar Rates of both light-limited and light-saturated photosynthesis and daily primary production were 2 to 6 times greater in the plankton than in the SIMCO. Bacterial growth rates ranges from 0.7 to 1.5 d^-1 in all three communities; however, because of the greater bacterial biomass in the plankton, bacterial production was 15 to 20 times higher there than in the SIMCO. These results suggest that during the early austral spring, planktonic production contributes significantly to total production in ice-covered environments.     

Inter-specific variation in anti-predator behavior in sympatric species of kangaroo rat

Kangaroo rats, Dipodomys, occupy desert habitats with little cover and thus are under high predation risk from diverse predators. The behavior used to assess predation risk or to escape capture is unknown. We therefore compared anti-predator behavior of two sympatric species of kangaroo rat of different sizes, D. merriami and D. spectabilis. We first examined whether kangaroo rats use olfaction as a first line of defense against snake predation and tested the rats for their responses to scent extracted from two species of snake that live sympatrically with the kangaroo rats, the Mojave rattlesnake (Crotalus scutulatus) and the gopher snake (Pitophis melanoleucus). We also tested for species differences in anti-predator behavior through 15-min interactions between the kangaroo rats and free-moving gopher snakes. We found that D. spectabilis actively approached the scent of both rattlesnakes and gopher snakes more than controls of vegetable oil and evaporated solvent (Fig. 1). In contrast, D. merriami did not differentiate snake odors from controls in the experimental arena, but they sniffed the sand where a free-moving snake had passed more than D. spectabilis. Both species successfully avoided predation in encounters with live snakes. Although total numbers of approaches and withdrawals were similar (Fig. 2), D. spectabilis spent significantly more time within striking distance of the snake than D. merriami. D. spectabilis approached the head of the snake in 93% of its approaches and often engaged in nose to snout contact with the snake. If the snake struck, D. spectabilis jumped directly backward to avoid a strike and footdrummed at a safe distance. In contrast, D. merriami oriented to the snake more than D. spectabilis, but approached the head in only 41% of the approaches and rarely engaged in nose-to-snout contact. The snakes struck, hissed and decreased predatory approaches with D. spectabilis but not with D. merriami (Fig. 3). These results show that kangaroo rats can behaviorally influence the risk of being preyed on by snakes. The two species differ, however, in how they react to snakes. The larger D. spectabilis confronts snakes while the smaller D. merriami monitors snakes from a safe distance and avoids them.     

宁波市某人工泻湖浮游生物变化特征及影响因素分析

以宁波市北仑区梅山水道形成的人工泻湖为研究对象,在不同季节进行水质及浮游生物调查,分析其浮游生物时空分布特征与水质的关系。4个采样点共检出浮游植物66种,以硅藻为绝对优势种,检出浮游动物25种,主要为桡足类、少量轮虫及网纹虫;拦坝后水道内浮游生物密度有了数量级增长,各项生物评价指数降低,但各采样点仍处于中污染水平。监测理化参数表明,研究水域在拦坝后盐度下降、悬浮物浓度下降,氮磷含量无明显变化;水域大部分点位处于中度富营养化水平。结合浮游生物分布与理化参数进行分析,发现堤坝合龙后,水道内侧海水淡化、悬浮物含量下降,导致浮游生物密度上升、生物多样性下降、出现淡水优势种群;营养盐含量不是浮游生物生长的限制因子,对浮游生物分布无显著影响。     

Phylogeography of the pipefish, <Emphasis Type="Italic">Urocampus carinirostris</Emphasis>, suggests secondary intergradation of ancient lineages

We assayed the pattern of mitochondrial DNA evolution in the live bearing, seagrass specialist pipefish, Urocampus carinirostris, in eastern Australia. These life history attributes were predicted to result in strong phylogeographic structure in U. carinirostris. Phylogenetic analysis of cytochrome b sequences detected two monophyletic mtDNA clades that differed by 8.69% sequence divergence - a large level of intraspecific divergence for a marine fish. The geographical distribution of clades was non-random and resembled clinal secondary intergradation over a 130-km stretch of coastline. Contrary to phylogeographic predictions, this large phylogeographic break does not occur across a traditionally recognised biogeographic boundary. Analyses of historical demography suggested that individuals belonging to the most widespread clade underwent a population expansion from a small refuge population during the Pleistocene.     

Feeding behavior of the Atlantic mackerel Scomber scombrus on the hydromedusa Aglantha digitale

Under laboratory conditions, the Atlantic mackerel Scomber scombrus feed on Aglantha digitale, a small, transparent medusa common in temperate and boreal waters. The same feeding response to A. digitale was observed whether the medusae were offered alone or together with a mixture of copepod prey. A. digitale mortality in the path of the mackerel school is estimated to be near 100%. Median carbon and nitrogen content of A. digitale was equivalent to about ten small copepods (e.g. Pseudocalanus sp.). A. digitale is abundant in temperate-boreal waters and may be an important source of nutrition to mackerel in the northwest Atlantic.     

Fine-scale distribution of larval fishes: patterns and processes adjacent to coral reefs in Kaneohe Bay,Hawaii

Plankton samples were taken from January to June 1987 in Kaneohe Bay, Oahu, Hawaiian Islands, with a free-fall plankton net, to investigate the fine-scale distribution of larval fishes around coral reefs. Daytime samples indicated that the postflexion larvae of two gobiids (Psilogobius mainlandi and an unidentified species) were significantly more abundant at stations immediately adjacent to reefs (near-reef) than at stations in open water off the reef (off-reef). These postflexion gobiid larvae appeared to be capable of resisting advection and dispersal while remaining in the water column near suitable adult habitats. The larvae of Foa brachygramma (Apogonidae) and Encrasicholina purpurea (Engraulidae) were significantly more abundant at off-reef stations than at near-reef stations. Nighttime samples indicated that the gobiid larvae depend on visual cues to remain near the reef. The horizontal distributions of F. brachygramma and E. purpurea larvae appeared to be related to their vertical positioning. These data suggest that typical ichthyoplankton surveys which do not sample close to adult fish habitats would greatly underestimate the abundances of larvae such as the gobiids.     

Vision in the rufus snake eel,<Emphasis Type="Italic"> Ophichthus rufus</Emphasis>: adaptive mechanisms for a burrowing life-style

A unique retinal topography was found in the rufus snake eel, Ophichthus rufus, a burrowing fish that shows a preference for nocturnal habits. The cells in the ganglion-cell layer formed a high-density semicircular band that covered the dorso-rostral, temporal and ventro-rostral areas of the retina. Three regions of high acuity were identified in the ventro-temporal, ventro-rostral and dorsal regions of the retina. This is the first time that multiple specialisations have been found in a benthic deep-water fish. The ventro-temporal and ventro-rostral visual axes were found at 48° and 54° from the retinal centre, subtending the dorsal field of view, while the dorsal axis was at 66°, subtending the ventral field of view. These characteristics may represent an adaptation to identify the mouth of the muddy burrows in which this species hides or where it looks for food. O. rufus has a duplex retina with numerous slender rod photoreceptors arranged in banks of four layers. This morphology is typical of deep-sea teleosts with a high visual sensitivity, which is useful in a dim-light environment. A few tiny cones were also present, probably enhancing acuity during the eel's daily activity. The retina of O. rufus also possessed a well developed vascularisation, possibly to overcome the hypoxic conditions that may be found in the burrow. The characteristics of the retina of the rufus snake eel indicate singular visual capabilities that are discussed in relation to the burrowing life-style and to what is known of feeding habits of this fish.     

Evaluation of fatty acids as biomarkers for a natural plankton community. A field study of a spring bloom and a post-bloom period off West Greenland

An investigation of the fatty acid composition of a natural arctic plankton community was carried out over two fishing banks located between 63°N and 65°N off the West Greenland coast. Samples for fatty acid analyses, species determination and biomass assessments of the plankton community were taken at the depth of fluorescence maximum. High biomass and diatom dominance during the spring bloom and low biomass and flagellate dominance in the post-bloom period were reflected by the fatty acid profiles. The total amount of fatty acid ranged from 55 to 132 µg l^-1 during the spring bloom and from 1 to 5 µg l^-1 during the post bloom. Analysis of the fatty acids showed that when the plankton was dominated by diatoms of the genera Thalassiosira and Chaetoceros, the proportions of C16:1(n-7) and C20:5(n-3) were correspondingly high. C18s, and particularly C18:1(n-9), were more abundant when the plankton was dominated by small autotrophic flagellates, primarily haptophytes. We found a good positive correlation between the common diatom marker, C16:1(n-7)/C16:0, and the biomass percentage of diatoms (r=0.742, P<0.001), as well as between the biomass percentage of flagellates and total C18 fatty acids (r=0.739, P<0.001). This supports the use of these specific fatty acids and fatty acid ratios as general biomarkers of the plankton community. However, the fatty acids are not specific enough to sufficiently characterise the composition of the plankton community, and microscopical support is needed to verify observed trends.     

Lipids and fatty acids as indicators of egg condition,larval feeding and maternal effects in Cape hakes (<Emphasis Type="Italic">Merluccius paradoxus</Emphasis> and <Emphasis Type="Italic">M. capensis</Emphasis>)

Cape hakes, Merluccius paradoxus and M. capensis, are important gadoid fish that are commercially harvested in the Benguela Current system off Namibia and South Africa. The aim of this study was to elucidate the nutritional condition and feeding preferences of their larvae. Hake eggs and larvae were sampled in austral spring of two consecutive years, 2007 and 2008, off the west coast of South Africa. They were identified to species using genetics, and total lipid content and fatty acid (FA) composition were analysed for each individual egg and larva to compare the condition of different early life stages of both hake species. Higher abundances of M. paradoxus eggs and larvae were consistently found compared to M. capensis. In both species, eggs contained wax esters (WE) and had significantly higher lipid content per dry mass than larvae. Lipid content as well as FA composition changed with the developmental stage of larvae. Quantities of essential fatty acid (EFA) increased with feeding of larvae due to dietary lipid incorporation. In 2007, yolk-sac larvae contained significantly lower total lipids than in 2008. It is argued that this was due to reduced lipid transfer by the spawning females to the eggs. These findings indicate that maternal effects are important in determining condition of hake larvae and that this may have an effect on their survival and subsequent recruitment.     

Diet composition and feeding intensity of horse mackerel,<Emphasis Type="Italic"> Trachurus trachurus</Emphasis> (Osteichthyes: Carangidae) in the eastern Adriatic

Diet composition and feeding intensity of the horse mackerel, Trachurus trachurus, collected in the eastern Adriatic Sea, were examined. Stomach contents of 1,200 specimens, taken at monthly intervals (January–December 1996), were analyzed. The stomachs contents consisted of five major prey groups: Crustacea (Euphausiacea, Mysidacea, Decapoda), Cephalopoda and Teleostei. Euphausiacea constituted the most important food resource by weight, number and frequency occurrence. Teleosts were the second most important food category, while mysids, decapods and cephalopods were occasional food. There was little seasonal variation in diet. Euphausiids were dominant prey during all seasons, and were especially abundant from spring and autumn. Feeding intensity of horse mackerel varied during the year. The lowest intensity of feeding was recorded in winter (February, March) and early spring (April) during the lower sea temperature as well as at the time of intensive spawning. Feeding activity increased upon spawning period (May and June) and was also higher during summer. Feeding intensity and diet composition changed during the diurnal cycle. The highest feeding intensity was recorded at night and during early morning hours. Euphausiids, mysids and a greater part of teleosts dominated night and morning diet, while decapods and cephalopods were most frequently in the daily and evening diet.Communicated by O. Kinne, Oldendorf/Luhe     

Operational sex ratios and behavioural sex differences in a pipefish population

In the pipefish Syngnathus typhle, only males brood embryos in specially developed brood pouches, supplying oxygen and nutrients. Laboratory studies have shown that this elaborate paternal care has led to sex-role reversal in this species: males limit female reproductive rate, females are the primary competitors for mates and males exercise greater selectivity in accepting mates. In the first field study of this pipefish, we describe mating behaviour in the wild and test the hypothesis that temporal variations in the operational sex ratio (OSR) determine sex differences in mating behaviour. Our study comprised two reproductive seasons of two sequential mating periods each, the latter separated by a lengthy interval of male brooding. During mating periods, females displayed to all males without wandering and males moved about searching for females, without reacting to all females. The OSR was least female-biased (or even male-biased) at the onset of the breeding season, when most pipefish were simultaneously available to mate, but became strikingly female-biased as males' pouches were filled. The OSR remained substantially female-biased during the second mating period, because few males became available to remate at any one time. As hypothesised, female-biased OSRs resulted in more female-female meetings. As well, females were above the eelgrass more often than brooding males, thus exposing themselves to conspecifics and/ or predators. In the second year, males arrived earlier than females on the breeding site and male pregnancies were shorter, because of higher water temperatures, so rematings occurred earlier. Males met more often during that year than the previous one, but male competitive interactions were still not observed. The field results support laboratory studies and demonstrate that behaviours associated with female-female competition are more prominent when the OSR is more female-biased. Correspondence to: A. Vincent