Predation of Northern Pike (Esox lucius L.) Eggs: A Possible Cause of Regionally Poor Recruitment in the Baltic Sea

Northern pike (Esox lucius) spawning habitat and egg mortality were studied in three spawning areas in 2001 along the Swedish coast of the Baltic proper: the Blekinge Archipelago, Kalmar Sound and coastal freshwater streams. Spawning peaked during the last week of April in streams, at temperatures ranging from 7.7 to 8.9 °C and during the first week of May in brackish waters, at temperatures ranging from 8.9 to 13.8 °C. Spawning occurred in shallow waters, at depths between 0.2–1.5 m, but generally most of the spawn was found in the shallowest areas. In streams, eggs were mainly attached to emersed vegetation, while in brackish sites pike eggs were well scattered among flooded emersed plants, submersed plants and filamentous algae. Mean egg density varied between 469–1829 m^–2 with the lowest density observed in Kalmar Sound. The calculated egg loss occurring from approximately one day after spawning to one day before hatching ranged from 41±7% in coastal streams to 67±6% in the Blekinge Archipelago and 100% in Kalmar Sound. The significant removal of eggs from spawning sites in Kalmar Sound and Blekinge was most likely due to predation from several fish species. In situ observations and stomach analyses suggested that many pike eggs in Kalmar Sound were lost to the three-spined stickleback (Gasterosteus aculeatus), a species that presently dominate the littoral small fish community. This study therefore suggests that egg predation by sticklebacks and other fishes may be a possible cause of the reported poor recruitment of coastal pike populations in the Kalmar Sound region.     

Reproductive behavior of largemouth triplefin Ucla xenogrammus (Tripterygiidae) on reefs of Kuchierabu-jima Island, southern Japan

Reproductive behaviors of the largemouth triplefin Ucla xenogrammus were observed on reefs of Kuchierabu-jima Island, southern Japan. Males exclusively maintained home ranges including spawning sites on vertical walls of overhanging reefs where filamentous algae were densely distributed. Females visited a male's home range to release adhesive eggs into the dense algae. Males guarded multiple egg clutches, but rarely showed fanning behaviors or mouth cleaning to the eggs. Aquarium experiments showed that guarding males had no significant effect on egg development. Because of the favorable water exchange conditions around their spawning sites, male U. xenogrammus could avoid the energy costs for egg care.     

Non-copulatory spawning and female participation during early egg care in a marine sculpinHemilepidotus gilberti

The reproductive behavior of a marine sculpin,Hemilepidotus gilberti, was observed in the aquarium. The behavioral sequence comprised fight for terriotory, courtship and fight for females, spawning and caring for the egg mass. Females very slowly deposited eggs enclosed with viscous ovarian fluid onto the substrate. Almost all the eggs contained in the ovary were released in a single spawning, its completion requiring 21–22 min. Territorial males emitted of spawning. Sneaking by neighboring males followed subsequently. This was the first observation of non-copulatory spawning among marine sculpins. Females cared for the eggs for severl days after spawning, and then male alone care continued until hatching (47–49 days after spawning). Participation by females in egg caring is likely to be caused by their spawning only once during the spawning season and the consequent lack of urgency for post-spawning feeding.     

Aspects of the spawning of western Atlantic butterflyfishes (Pisces: Chaetodontidae)

Synopsis The status of knowledge of spawning among the five shallow waterChaetodon species in the western Atlantic is reviewed. Spawning has been observed for three species in Puerto Rico, St. Croix and the Bahamas, with possible courtship in a fourth.Chaetodon aculeatus spawned near the time of sunset over objects on the reef as single female/male pairs or as two females and one male, with pair spawning in rapid succession. Spawning occurred during much of the lunar month from February to April and it is uncertain whether any lunar periodicity to spawning exists. Male-male aggression was noted. Spawning sites (coral heads) were alternated daily and it is likely that females spawn only once every two days. A single female produced as many as 2090 eggs in a single spawning.Chaetodon capistratus spawned during much of the lunar month from February to April. It spawned about 5 min afterC. aculeatus, occasionally using the same sites, and alternated sites daily. A female produced as many as 3710 eggs in one spawning.Chaetodon striatus spawned from February to April but it is unknown if it has any lunar spawning cycle. No predation attempts by piscivores on spawning adults were seen. Predation byMelichthys niger on eggs ofC. striatus occurred. No egg predation was observed forC. aculeatus andC. capistratus. With an assumed four month reproductive season, alternate day spawning and observed egg production values,C. aculeatus andC. capistratus produce respectively about 100 000 and 200 000 eggs per large female per year. The reproductive strategy of smaller species may be to produce moderate numbers of eggs per day over a spawning season of at least a few months while larger species may produce more eggs per day for a shorter period.     

Lek-like spawning,parental care and mating periodicity of the triggerfishPseudobalistes flavimarginatus (Balistidae)

Synopsis The triggerfishPseudobalistes flavimarginatus has a lek-like spawning system at Yonge Reef, Great Barrier Reef. Males migrate to a traditional mating ground where they establish territories enclosing nest sites and egg chambers. Females arrive several days later and choose a male for mating. There is biparental care of the fertilized eggs and a strict division of labour. Females appeared to be confined to the nest site by the male. Mating was semi-lunar: eggs were observed in nests several days before the new and full moons on days when high tide occurred near sunset. Differences were observed in the spawning system of another triggerfish (P. fuscus) between the Great Barrier Reef and the Red Sea. Observations of another triggerfish (Balistoides viridescens) are also reported.     

Embryonic development and reproductive seasonality of Buccinanops globulosus (Nassariidae) (Kiener, 1834) in Patagonia,Argentina

Buccinanops globulosus mated all year round, with higher frequency from May to September, prior to spawning months. Gravid females were found between October and March. Oviposition peaked during rising temperatures and longest daylength while hatching peaked with high water temperature and declining daylength. Gravid females measured between 20 and 41?mm in shell length. The spawn consisted on average of 31 egg capsules, each containing 1266 eggs. Embryos usually completed development within each egg capsule by ingesting small fragments of the uncleaved nurse eggs, which were not a limiting resource. Egg capsules with more than one embryo were not common; in those cases, the embryos had different sizes probably related to intracapsular competition for nutrients, and were on average smaller than solitary embryos in the other capsules. Embryos hatched as crawling juveniles with a mean hatchling shell length of ~3.4?mm. In a few cases, malformed embryos were found, but it was not a common phenomenon. The information recorded in this study, as the minimum reproductive size and spawning season, is valuable for fisheries management.     

Nonselective use of vegetation for spawning by the diadromous fish Galaxias maculatus

Terrestrial egg development is advantageous for the amphidromous fish Galaxias maculatus because it increases access to oxygen, increases incubation temperatures, and reduces aquatic predation. The characteristics of New Zealand's riparian vegetation have changed considerably since colonial times from native vegetation to exotic grasses, with potential effects on the spawning dynamics of fish. Here, we used a series of experiments to test preferences of G. maculatus for egg laying in combinations of a native sedge and rush, and two exotic grasses. In laboratory experiments, G. maculatus spawned in all available habitats with no preference for native over exotic vegetation. Egg survival did not differ between the vegetative habitats, but in a treatment where only river stones were available, eggs were laid but they dehydrated and died. In a field experiment, there was no evidence for a preference for spawning in native vegetation, and no association between a “key” feature of vegetation—tiller density—and the number of eggs laid. The microclimate beneath dense vegetation in experimental treatments was more constant than ambient conditions with few extreme temperatures and low humidities. Given the readiness of G. maculatus to spawn in exotic vegetation, there appears to be no benefit in restoring native vegetation to enhance egg laying or survival of these taxa.     

Egg-feeding in the freshwater piscicolid leech Cystobranchus virginicus (Annelida, Hirudinea)

Abstract. Leeches are common terrestrial and aquatic annelids, parasitizing or feeding on a wide range of host taxa. Here we report evidence for an unusual feeding behavior of egg‐feeding in the piscicolid leech Cystobranchus virginicus. We identified distended specimens of C. virginicus in the nests of at least 4 different fish species: Campostoma anomalum, Moxostoma carinatum, Moxostoma sp. (either M. anisurum and/or M. breviceps), and Nocomis leptocephalus. We collected a total of 41 leeches from the nests of these host species and documented at least 1 leech in 19 of 55 nests (35%), with many sites containing multiple leeches. Individuals of C. virginicus were not identified feeding on any of the 41 adult specimens of Moxostoma spp. or the 635 adult specimens of Nocomis leptocephalus examined, and were never found in the absence of active host spawning (26 sites). These results are consistent with individuals of C. virginicus being an opportunistic or possibly even an obligate egg‐feeder, potentially timing their own reproductive activities with the spawning of their fish hosts. The current distribution of C. virginicus has been expanded to include North Carolina, South Carolina, and Tennessee. The potential for leech species to induce mortality in developing fish eggs could be a concern for fish conservation and merits further investigation.     

Reproductive behavior of the Atlantic silverside,Menidia menidia (Pisces,Atherinidae)

Synopsis The spawning behavior of the Atlantic silverside,Menidia menidia, was studied at two sites on the North Edisto River estuary in South Carolina. Prespawning schools moved back and forth along the shoreline as the time of high tide approached. Spawning runs took place in the upper intertidal zone at high tide.Spawning fish deposited their eggs on three types of substrates: 1) the lower stems of cordgrass plants,Spartina alterniflora, 2) detrital mats, and 3) exposed cordgrass roots along erosional scraps. Spawning behavior during egg deposition and fertilization was similar for all three substrates. Females released eggs during a rapid fluttering motion of the posterior half of the body. A similar movement accompanied release of sperm by males. Behavior of fish just prior to spawning insured deposition of gametes at locations that provided protection from thermal and drying stress during development. Eggs were deposited at mean intertidal elevations of 1.8 and 1.5 meters above mean low water (MLW) at respective study sites. They were exposed to the atmosphere for approximately ten hours between successive high tides.During spawning runs in which eggs were deposited at the base of cordgrass plants, ambient dissolved oxygen concentrations of the water in the spawning area were sometimes reduced to < 1.0 mg. 1^–1. Spent fish, apparently incurring an oxygen debt while spawning, formed a nonschooling aggregation offshore from the spawning zone.The recurrent use of specific spawning substrates for egg deposition resulted in an uneven distribution of spawning runs along the shoreline at each study site.Contribution No. 409 to the Gulf Breeze Environmental Research Laboratory. Contribution No. 377 to the Belle W. Baruch Institute for Marine Biology and Coastal Research.     

Spawning and subsequent copulating behavior of the elkhorn sculpinAlcichthys alcicornis in an aquarium

The spawning and subsequent copulating behavior of the elkhorn sculpin (Alcichthys alcicornis) were observed in an aquarium. Soon after the female was introduced into the tank, the male approached and initiated courtship behavior by unfolding all fins. Females which carried ovulated eggs eventually accepted the male and spawning progressed. Females which had not ovulated, repeatedly rejected the male’s advances. The onset of spawning in the female was indicated by fluttering of the pectoral fins and successive horizontal undulations of the posterior trunk which served to sweep the spawning site. The laying of several thousand eggs was preceded by the female taking several large inhalations of water through the mouth and a slight raising of the posterior trunk. The male continued various behavior related to courtship while the female approached spawning. After spawning, the female smoothed the egg mass into a thin layer by using the posterior trunk and anal fin. After spawning, and while the female flattened the egg mass, the male copulated repeatedly. No copulation was observed prior to spawning. Semen was observed to be emitted from the tip of the penis and to leak from the female’s genital pore. Eggs were fertilized externally by the emitted sperm in the case of females which had no prior copulating experience in the present breeding season. The entire reproductive behavior of the pair ceased, as the female left the spawning site after flattening the egg mass. The male remained near by, but did not aerate the eggs. During the breeding season, the area around the female’s genital pore and along both sides near the base of the anal fin became slightly distended and soft by the accumulation of a serous fluid. It is presumed that this edematous modification serves to flatten the egg mass for softness without crushing the eggs.     

Spawning strategy in Atlantic bobtail squid <Emphasis Type="Italic">Sepiola atlantica</Emphasis> (Cephalopoda: Sepiolidae)

This study aimed to determine the spawning strategy in the Atlantic bobtail squid Sepiola atlantica, in order to add new information to the knowledge of its reproductive strategy. A total of 12 females that spawned in aquaria were examined. Characteristics of the reproductive traits and egg clutches were similar to those of other known Sepiolidae. Clutch size varied from 31 up to 115 eggs. Females of this species had incorporated up to 1.58 times of their body weight into laid eggs. The size of laid eggs showed a positive correlation with maternal body size, supporting the idea that female size is a determinant of egg size. Our data suggest that S. atlantica is an intermittent terminal spawner, and that its spawning strategy comprises group-synchronous ovary maturation, multiple egg laying, and deposition of egg clutches in different locations. The obtained data provide insights for future comparative studies on reproductive allocation.     

Spawning habitat selection and suitability for Japanese dace, Tribolodon hakonensis

We have investigated the effects of three current velocities and three substrate sizes (gravel to cobble) on the spawning behavior of Japanese dace, Tribolodon hakonensis, in spawning-induction and habitat-selection experiments. In the spawning-induction experiment under laboratory conditions the number of females induced to spawn was significantly fewer at low current velocity (ca. 5 cm s^–1) than at medium (ca. 30 cm s^–1) or high (ca. 50 cm s^–1) current velocity. Females spawned independently of substrate size in the experimental tank, and did not bury their eggs in the substrate bed. In the habitat-selection experiment under field conditions, females selected the substrate microhabitat and spawned more frequently at a site with medium substrate size (very coarse gravel; major axis ca. 40 mm) than at sites with small (fine gravel; ca. 8 mm) or large (cobble; ca. 70 mm) substrate size. We conclude that Japanese dace select spawning sites at least partly on the basis of current velocity and substrate size, which affects the survival rate of eggs.     

Reproduction in the giant clams Tridacna gigas and T. derasa in situ on the north-central Great Barrier Reef,Australia, and Papua New Guinea

Small groups of T. gigas and a single individual of T. derasa have been observed to spawn sperm in nature during the austral summer on the Great Barrier Reef. In T. gigas diel periodicity to the spawning generally coincided with incoming tides near second (full), third and fourth (new) quarters of the moon phase. Intense spawning ranged from 30 min to 2 1/2 h, with spawning contractions occurring every 2–3 min. Eggs were never observed being spawned in nature. A new technique, an egg catching device, was used for sampling eggs of T. gigas which were observed spawning sperm. Results indicated that the egg phase of spawning did not follow spawning of sperm. Some clams which do not respond to a spawning neighbour may be reproductively inactive. In 70% of observations of nearest spawning neighbours the clams were within 9 m of one another. A hypodermic extraction technique was used for clam gonadal material. Examination of tissue suggested a potential early to mid-austral summer spawning for T. gigas, and to a lesser degree for T. derasa.     

Host handling behavior of the egg parasitoid Telenomus triptus to the egg mass of the stink bug Piezodorus hybneri

Host handling behavior of Telenomus triptus Nixon (Hymenoptera: Scelionidae) to an egg mass of Piezodorus hybneri Gmelin (Heteroptera: Pentatomidae) was studied in the laboratory. Five distinct behavioral events could be distinguished in the host handling behavior: drumming, ovipositor-insertion, marking, walking, and resting. Female wasps showed two types of behavioral sequence in an ovipositional bout. One consisted of drumming, ovipositor-insertion, and marking, and the other drumming, ovipositor-insertion, walking, and resting. Females did not seem to lay an egg when ovipositor-insertion was not followed by marking. This was observed frequently in the early oviposition bout, on average 2.9 times per female. Females finally succeeded in parasitizing all the eggs in a host egg mass in most cases. The durations of drumming, ovipositor-insertion, and marking on an egg mass were nearly constant, while the total time spent by a female on and beside a host egg mass varied considerably as a result of variable durations of walking and resting. Self-superparasitism occurred when almost all the eggs in a host egg mass were parasitized. Females laid the first male egg within the first four eggs; this could be an adaptation to small egg masses or single egg.     

Spawning timing of the cleaner wrasse, Labroides dimidiatus, on a warm temperate rocky shore

We observed spawning behaviors of the haremic cleaner wrasse Labroides dimidiatus at a rocky reef in southern Japan. Females released pelagic eggs daily from June to September after a spawning ascent by pairs. When the high tide occurred between noon and evening (quarter moon to new or full moon), they spawned around the time of the tide providing fast offshore currents. For the rest of the lunar cycle, spawnings occurred during late afternoon independent of the state of the tide at the time of day. This spawning pattern is quite different from that of conspecifics on coral reefs that always spawn around the high tide at all lunar stages to avoid potential egg predators. The wrasse ascended high in the water column, probably because of its predator-immune characters as a cleaner. The steep slope of the study site seemed to contribute to releasing gametes far above aggregating planktivores. Thus, gametes were not subject to heavy predation. Current direction might not always affect survival of spawned pelagic eggs on the temperate reefs so crucially as it does on coral reefs. We conclude that spawning during unfavorable tidal conditions during the daytime may be better than spawning during the best tidal condition in the crepuscular period when predation pressure on adults will be high. Intraspecific variation in the timing and location of spawning of the wrasse may result from different conditions for larval survival. Received: February 6, 2000 / Revised: June 16, 2000 / Accepted: July 17, 2000     

Spatial aspects of reproduction in buffalo sculpin,Enophrys hison

Synopsis The spatial distribution and parental care of demersal egg masses of buffalo sculpin (Enophrys bison) were studied at 11 areas in Puget Sound, Washington, during 1972–1973. Adult buffalo sculpin inhabited rock and rock-sand habitats at depths from the intertidal to minus 20 m throughout the year, but 84% of all spawn clusters were found at depths between –1 m and +1 m within localized rocky areas. Areas used for spawning were exposed to strong tidal currents and other water movements. A single male fish guarded and fanned spawn, but eggs were temporarily deserted when less than about 1 m of water covered the spawning site. The cyclical nature of male desertion, behavior of spawn predators, and spawn mortality due to physical factors while emergent suggest several possible advantages to low intertidal spawning. These advantages include spatial and temporal escape from marine predators by embryos, time for the tending parent to forage, and optimal physical conditions for embryonic development.     

Reproductive behaviour of the filefishRudarius ercodes: male spawning parades and female choice

Synopsis The reproductive behaviour ofRudarius ercodes is described from undersea observations in Aburatsubo Bay, Japan. Reproductive behaviour can be separated into four parts: (1) Prespawning Search = searching for spawning sites by females and searching for gravid females by males, (2) Spawning Parade = males follow a gravid female in a line, competing with each other to reach the head of the queue, (3) Spawning = the female takes the spawning position, males rush to the side of the female, and mating occurs between one female and several anterior males of the spawning parade, (4) Parental Care = females attach adhesive eggs to seaweed with the mouth and guard them until embryos hatch. There is no male parental care. The reproductive season ranges from May to October and spawning occurs early in the morning every day. Females begin feeding early in the morning, but males feed little at this time.R. ercodes shows neither territorial behaviour nor fixed-pair spawning. One male might spawn several times in one morning. One female spawns at most once every 5 days. The mating system of this species is promiscuous. The probable function of the spawning parade as a style allowing female choice is discussed.     

Sound Production in Codoma ornata (Girard) (Cyprinidae)

Male Codoma ornata produce sounds associated with aggression and spawning activities during the breeding season. Females do not produce sounds. Males most often produced sounds associated with escalated displays of aggression, courtship and the spawning act. C. ornata spawn in crevices, but previously were reported to spawn as egg-clusterers in cavities. Structurally, sounds are low frequency, vary in duration according to context and are not harmonic. The mechanism of sound production is unknown.     

Timing of paddlefish spawning in the Upper Missouri River,Montana, USA in relation to river conditions

Spawning activity of paddlefish Polyodon spathula in the Missouri River, Montana in 2008–2009 was examined to delineate spawning sites and times in relation to discharge, water temperature and turbidity. One hundred thirty‐six eggs were collected at water temperatures ranging from 12.0 to 20.7°C (mean, 16.3°C; SD, 2.5). Only 12 of 89 (13%) congregations of radio‐tagged adults observed during the spawning period coincided with egg captures. Six larvae were collected at water temperatures ranging from 19.1 to 21.7°C (mean, 20.5°C; SD, 0.86). Peak discharge in 2008 (903 m³ s^?1 on 14 June) was approximately 30% greater in magnitude and occurred 11 days later than peak discharge in 2009 (612 m³ s^?1 on 3 June). Despite these differences in the hydrograph, no significant differences in egg CPUE were found between years (anova , F = 0.69, P = 0.56). Logistic regression identified no significant river condition variables associated with the presence or absence of eggs (P > 0.14 for all variables). However, in both years maximum egg CPUE was recorded within 3 days of the hydrograph peak and at similar water temperatures (17.5°C in 2008, 16.8°C in 2009). These results suggest an overall association of peaking discharge and seasonally warming water temperatures with egg deposition. Higher catches of eggs and larvae than observed in this study may be necessary to clarify short‐term (day‐to‐day) effects of environmental changes on spawning activity. Continued investigation of the relationship between short‐term changes in river conditions and paddlefish spawning activity is needed to understand the mechanics underlying the reproductive success of this species.     

Evidence of autumn spawning in Suwannee River Gulf sturgeon,Acipenser oxyrinchus desotoi (Vladykov, 1955)

Evidence of autumn spawning of Gulf sturgeon Acipenser oxyrinchus desotoi in the Suwannee River, Florida, was compiled from multiple investigations between 1986 and 2008. Gulf sturgeon are known from egg collections to spawn in the springtime months following immigration into rivers. Evidence of autumn spawning includes multiple captures of sturgeon in September through early November that were ripe (late‐development ova; motile sperm) or exhibited just‐spawned characteristics, telemetry of fish that made >175 river kilometer upstream excursions to the spawning grounds in September–October, and the capture of a 9.3 cm TL age‐0 Gulf sturgeon on 29 November 2000 (which would have been spawned in late September 2000). Analysis of age‐at‐length data indicates that ca. 20% of the Suwannee River Gulf sturgeon population may be attributable to autumn spawning. However, with the very low sampling effort expended, eggs or early life stages have not yet been captured in the autumn, which would be the conclusive proof of autumn spawning. More sampling, and sampling at previously unknown sites frequented by acoustic telemetry fish, would be required to find eggs.