Beach Spawning in Fishes: Phylogenetic Tests of Hypotheses

Marine fishes that spawn at the water's edge or even out ofwater provide their eggs with the advantages of the warmer temperaturesand high oxygen availability of the high intertidal zone. However,they increase the risks of desiccation and terrestrial predation.Beach spawners are present in at least 6 families of teleostfishes. Two hypotheses about the origin of beach spawning aretested by mapping of reproductive habitat and spawning siteutilization onto phylogenies of two families that contain multiplespecies that spawn on beaches: Osmeridae, the smelts, and Atherinopsidae,the silversides. Our analysis suggests that beach spawning hasevolved repeatedly in certain lineages, and that its antecedentsare different for each family. Anadromy appears to have beenthe precursor for at least 3 different evolutionary originsof beach spawning in osmerids, while nearshore marine spawningin association with plant or gravel substrates was probablythe precursor for the atherinopsids. Phylogenetic analysis enablesus to reject or support specific evolutionary hypotheses foreach clade.     

Mass spawning of Caesio teres (Pisces: Caesionidae) at Enewetak Atoll,Marshall Islands

Synopsis Mass spawning of Caesio teres occurred between March and August, 1983 on a reef emerging from deep water just inside the East Channel of Enewetak Atoll, Marshall Islands. Aggregations and spawning were observed from one day before until three days after the full moon. Spawning aggregations were not observed during the new or quarter moons. Spawning commenced after high tide, when the current began to flow out the pass from lagoon to ocean. The spawning aggregation of close to 1000 individuals migrated to the spawning site. Spawning occurred when the aggregate ascended to near the water's surface. Subgroups dashed horizontally within the aggregate, releasing a highly visible gamete cloud. Predation on spawning adults was not observed. Predation on spawned eggs was noted. The eggs of C. teres are described. Mid-Pacific Research Laboratory, Enewetak Atoll, Republic of the Marshall Islands Present address: Motupore Island Research Station, P.O. Box 320, University of Papua New Guinea, Papua New Guinea     

Fish, Flows and Flood Plains: Links between Freshwater Fishes and their Environment in the Murray-Darling River System, Australia

Knowledge of the biology of native fishes of the Murray-Darling Basin is based largely on studies conducted under hatchery conditions and on a limited number of recreationally important species. From observations that increases in water level in aquaculture ponds initiate spawning in some species, and from limited studies of wild fishes and studies in overseas floodplain river systems, a perception has emerged of the importance of flooding and the flood plain in the life cycles of Murray-Darling fishes in general. However, there is little confirmatory evidence of the use of temporary floodplain habitats by larvae, juveniles or adults. The significance of in-channel habitats, especially for rearing, has received little attention. Murray-Darling fish species can be placed into three life history modes, based mainly on spawning style and time and developmental intervals of larvae at first feeding. Fish in each group may be able to take advantage of floods if the timing is right and prey are plentiful, however, the larvae of some species are able to recruit under non-flood conditions within the main river channel. This forms the basis of the low flow recruitment hypothesis, which attempts to explain why some species spawn during the warmest months and lowest flows and how they are able to recruit under these conditions. This hypothesis is then placed in the context of the current state of knowledge of the relationships between flow and the biology of Murray-Darling fishes, specifically cues for spawning, movement and recruitment. The lack of widespread evidence for floodplain use by any life history interval of fish may be due to a paucity of study, however, there are some fundamental factors, such as the predictability of timing and duration of high flow events as well as the lack of coincidence of high flows and high temperatures in some regions of the Basin, which may be important in determining the use of floodplain habitats by fish.     

兴修水利枢纽后汉江产漂流性卵鱼类的繁殖生态

汉江产漂流性卵的鱼类至少有25种,其中经济鱼类有草鱼、青鱼、鲢、鳙、鯮、鳤、鳡、长春鳊、赤眼鳟、铜鱼、吻鮈、鳜以及三种红鲌属鱼类。       

Population variation in spawning current speed selection in the blacktail shiner,Cyprinella venusta (Pisces: Cyprinidae)

Synopsis Many stream fishes lay demersal eggs. Some species simply scatter the eggs across the substrate, but many place the eggs in protected sites such as crevices (e.g., Cyprinidae:Cyprinella). Eggs laid in crevices may experience increased development times or lower embryo survival due to reduced water exchange rates compared to more open egg deposition sites. If so, the presence of an appreciable current near the crevice may be important for insuring water exchange. However, currents that are too strong may prevent complete insemination or prevent the eggs from adhering to the substratum before they are washed from the crevice. Thus, current speed preferences for spawning may be under strong stabilizing selection. Representatives of four populations of a crevice spawning stream fish (blacktail shiner,Cyprinella venusta), representing environments ranging from a swift stream to a lake, were studied in stream tanks to evaluate female spawning current use. The observed frequency of use of spawning currents was significantly different from random in all populations. Low current speeds were generally avoided, and a speed of ca. 30 cm sec^–1 was used most often. However, significant differences were observed among populations, and the variation correlated well with the apparent range of current speeds found in the four environments. Although other possible explanations cannot be dismissed, the results are consistent with the hypothesis that the populations have become adapted to their local current regimes.Authorship in order of seniority     

葛洲坝水利枢纽兴建后对青、草、鲢、鳙繁殖生态效应的研究

本文分析了葛洲坝枢纽截流后,大坝上、下游草、青、鲢、鳙的繁殖条件,产卵场位置,产卵规模,鱼卵、鱼苗成色以及4种鱼的群体组成等。指出除原宜昌产卵场发生变化外,其余产卵场基本存在,并在上游新发现了几处产卵场。讨论了大坝对4种鱼的影响程度和上游存在草鱼地方性群体,论证了4种鱼不必过坝产卵的理由。       

Tolerance of forced air emergence by a fish with a broad vertical distribution, the rockpool blenny, Hypsoblennius gilberti (Blenniidae)

Many intertidal fishes, particularly among the Blenniidae and Cottidae, possess amphibious adaptations, including the ability to breathe in air and to avoid desiccation in terrestrial conditions. These traits are absent in subtidal species of blennies and cottids. Hypsoblennius gilberti, the rockpool blenny, is found in shallow rockpools in the mid to high intertidal areas of Southern California, and deeper to 18 m in the subtidal zone. This broad vertical distribution could indicate that this blenny is adapted for tidal air emergence, although H. gilberti has not been observed out of water in its natural habitat. H. gilberti does not emerge voluntarily from hypoxic sea water in the laboratory, but it easily withstands 3 h out of water. The aerial respiratory exchange ratio (CO2 released compared to O2 consumed) is 0.70, similar to that of amphibious intertidal fishes in air, indicating sufficient release of metabolically produced CO2 while emerged. There is no increase in aquatic respiration following emergence. However, unlike other amphibious fishes that maintain aerial oxygen consumption at a level similar to aquatic oxygen consumption, H. gilberti has an aerial oxygen consumption rate one-third that in water. H. gilberti can recover rapidly from terrestrial water loss, and shows no change in evaporative water loss rates at 93% and 77% relative humidities. The amphibious capabilities in H. gilberti, even if rarely used, permit survival in air during tidal emergence. These findings suggest that H. gilberti may demonstrate an intermediate condition between the amphibious species of intertidal fishes that regularly emerge from water, and the subtidal fishes that do not survive air emergence and are completely restricted to an aquatic habitat.     

Observations of the migration of adult Dasylepida ishigakiensis (Coleoptera: Scarabaeidae) from belowground to the soil surface and their reproductive status by MRI

Field observations indicate that white grub beetles Dasylepida ishigakiensis Niijima et Kinoshita (Coleoptera: Scarabaeidae) pupate approximately 40 cm underground and then emerge from the soil surface as adults. Although the factors influencing adult emergence from the soil have been studied, the migration of adults from belowground to the surface has never been characterized. In this study, the vertical movement of virgin adults from natural eclosion depths to their emergence from the soil surface was investigated. Virgin adults were introduced at a depth of 40 cm from the soil surface in wooden-framed glass cases, and the vertical position of each individual was checked every day. Both females and males emerged near the soil surface at approximately day 50 after they were transferred from 24 °C to 18 °C conditions. The maturity of the ovaries in unemerged and emerged females was also observed by magnetic resonance imaging. All females that had emerged from the soil possessed developed oocytes, and most possessed mature eggs. The cross-sectional area of eggs was determined in magnetic resonance images and compared between unemerged and emerged females, and the egg areas of emerged females were significantly larger than those of unemerged females. Our results indicate that the timing of female emergence from soil corresponded to ovary maturity in this species.     

Features of reproduction of whitefishes

Based on features of reproduction, whitefishes (Coregonidae) differ from salmonids (Salmonidae) and graylings (Thymallidae). They differ from other salmonid fishes (Salmoniformes) in the following characteristics: they spawn, as a rule, during fall and winter; they do not construct spawning redds; their eggs are not protected; and substantial egg mortality is compensated by a high fecundity. The eggs of several whitefish species can develop in the conditions of “pagon” (within the ice). The main reasons for egg mortality at riverine spawning grounds are as follows: abnormalities during fertilization and development of the embryo, predation on the eggs, parasitic deceases, freezing of spawning grounds, and drift of the eggs from the spawning grounds. In addition (in the lakes), egg mortality can be caused by a deficit of oxygen, siltation, acidulation, and pollution of water. The survival of eggs of whitefishes in the Ural tributaries of the Lower Ob River in different years ranges from 0 to 93%. At a stable environment, the age of sexual maturation is connected with a certain body size of the fish. In the northern water bodies, the cycle of gonadal development can reach more than 1 year. Pollution of water by heavy metals, oil, and toxicants leads to serious anomalies of the reproductive system and to decreasing reproductive potential of the stocks.     

Invertebrate infestation on eggs and hatchlings of the loggerhead turtle, Caretta caretta, in Dalaman, Turkey

The damage caused by some invertebrates to the eggs and hatchlings of loggerhead turtles, Caretta caretta, was investigated during the summer of 2002 on Dalaman beach, Turkey. The specimens, identified to family or genus levels, from nine families representing seven orders were recorded as infesting nests of loggerhead turtles. The heaviest impacts on loggerhead turtle nests was made by Pimelia sp. (Tenebrionidae, Coleoptera). Twenty-four (36.3%) out of 66 intact loggerhead hatched nests were affected by these larvae. Larval damage by Pimelia sp. was recorded in 188 (10.6%) out of 1773 eggs, but only in two (0.28%) hatchlings. The results show that fewer insects were in the nest the further from vegetation and therefore the relocation of nests from the water's edge to further inland close to vegetation may increase the infestation rate of the eggs.     

The beach‐spawning California grunion Leuresthes tenuis eats and digests conspecific eggs

Apparent egg cannibalism was investigated in the beach‐spawning California grunion Leuresthes tenuis. Three hypotheses were tested to determine whether L. tenuis regularly consumes and efficiently digests conspecific eggs. First, examination of the gut contents of adults collected at four spawning sites over two seasons showed that the intestines of most fish from all the sites (57–87%, n ≥ 30, each site) contained L. tenuis eggs. The two other hypotheses focused on digestion of the eggs. First, the force required to crush cannibalized eggs was significantly less than that for uncannibalized eggs (fertilized or unfertilized), indicating that ingestion weakens the egg chorions. Second, conspecific eggs fed to fish held in the laboratory visibly degraded as they passed through the gut. The eggs lost c. half of their protein content and about two‐thirds of their lipid content as they passed from proximal to distal regions of the gut, indicating that digestion occurred. Digestive enzyme activities of the gut further confirmed that L. tenuis can break down the contents of ingested eggs. Trypsin activity decreased and aminopeptidase activity increased posteriorly along the gut, whereas amylase and lipase activities exhibited less clear patterns by gut region. As far as is known, this study is the first to show that L. tenuis is an egg cannibal.     

Metabolism,gas exchange,and acid‐base balance of giant salamanders

The giant salamanders are aquatic and paedomorphic urodeles including the genera Andrias and Cryptobranchus (Cryptobranchidae), Amphiuma (Amphiumidae), Siren (Sirenidae), and Necturus (Proteidae, of which only N. maculosus is considered ‘a giant'). Species in the genera Cryptobranchus and Necturus are considered aquatic salamanders well adapted for breathing water, poorly adapted for breathing air, and with limited abilities to compensate acid‐base disturbances. As such, they are water‐breathing animals with a somewhat fish‐like respiratory and acid‐base physiology, whose habitat selection is limited to waters that do not typically become hypoxic or hypercarbic (although this assertion has been questioned for N. maculosus). Siren and Amphiuma species, by contrast, are dependent upon air‐breathing, have excellent lungs, inefficient (Siren) or no (Amphiuma) gills, and are obligate air‐breathers with an acid‐base status more similar to that of terrestrial tetrapods. As such, they can be considered to be air‐breathing animals that live in water. Their response to the aquatic hypercarbia that they often encounter is to maintain intracellular pH (pH_i) and abandon extracellular pH regulation, a process that has been referred to as preferential pH_i regulation. The acid‐base status of some present‐day tropical air‐breathing fishes, and of Siren and Amphiuma, suggests that the acid‐base transition from a low PCO₂‐low [] system typical of water‐breathing fishes to the high PCO₂‐high [] systems of terrestrial tetrapods may have been completed before emergence onto land, and likely occurred in habitats that were typically both hypoxic and hypercarbic.     

Maturation, fecundity, and intertidal spawning of Pacific sand lance in the northern Gulf of Alaska

Pacific sand lance Ammodytes hexapterus in Kachemak Bay, Alaska, showed no sexual dimorphism in length-to-weight (gonad-free) ratio or length-at-age relationship. Most matured in their second year, males earlier in the season than females, but females (31%) attained a higher gonadosomatic index than males (21%). Sand lance spawned intertidally once each year in late September and October on fine gravel or sandy beaches soon after the seasonal peak in water temperatures. Sand lance in Cook Inlet and Prince William Sound displayed similar maturation schedules. Schools were dominated 2: 1 by males as they approached the intertidal zone at a site where spawning has taken place for decades. Sand lance spawned vigorously in dense formations, leaving scoured pits in beach sediments. Fecundity of females (93–199 mm) was proportional to length, ranging from 1468 to 16 081 ova per female. About half of the overall spawning school fecundity was derived from age group 1 females (55% of the school by number). Spawned eggs were 1·02 mm in diameter, demersal, slightly adhesive, and deposited in the intertidal just below the waterline. Sand lance embryos developed over 67 days through periods of intertidal exposure and sub-freezing air temperatures.     

Quantitative observations of predation during spawning rushes of the labrid fishThalassoma cupido at Miyake-jima,Japan

A rapid upward spawning rush is a nearly universal phenomenon among reef fishes that spawn pelagic eggs. Although spawning rushes have long been considered to have evolved as a defense against egg predators and/or attacks on the spawning fishes by piscivores, these hypotheses have never been tested in the field. We analysed piscivore attacks during three motor patterns associated with group spawning of the labrid fishThalassoma cupido at Miyake-jima, Izu Islands, Japan. Egg predation on group spawnings was also quantified. Of 206 piscivore attacks on spawning fishes by seven predator species, 17.1 (83.1%) occurred during 461 spawning rushes (1/2.7 spawning rushes). No attacks were successful during spawning rushes, and only four kills were made in 206 attacks summed from all three motor patterns associated with spawning, amounting to a piscivore success rate of only 1.9 %. In contrast, gametes from 90 of 213 spawnings (42.3 %) were consumed by nine species of planktivorous fishes. Spawning fish seemed not to recognize egg predators and made no attempt to avoid them, often spawning in the midst of waiting aggregations of damselfishes. Our data indicate that the rapidly of the spawning rush ofT. cupido serves as an excellent defense against piscivores, but is ineffective against egg predators. Evidence is presented from observations of seven other species suggesting that spawning ascents provide little protection against water column egg predators in predator-rich environments.     

Aerial release of CO2 and respiratory exchange ratio in intertidal fishes out of water

Synopsis Many fishes in the marine intertidal zone have the ability to exchange respiratory gases in air during brief periods out of water. Previous studies on marine amphibious fishes such as mudskippers and rockskippers have shown that they release CO₂ in air at a rate consistent with its release in aquatic respiration. Respiratory Exchange Ratios (RER, CO₂ released per O₂ consumed) are between 0.7 and 1.0, similar to Respiratory Quotients (CO₂ produced metabolically per O₂ consumed). However, previous studies of temperate intertidal fishes emerged only at low tides have been less consistent, with two species reported similar in ability to the amphibious skipper fishes, and two others reported with much lower RERs in air. This study examines 12 species of fishes, from six families (Batrachoididae, Cottidae, Gobiesocidae, Kyphosidae, Pholididae, and Stichaeidae), present in the intertidal zone of California. All 12 species exchanged O₂ and CO₂ in air. Like the amphibious skipper fishes of marine tropical waters, many intertidal fishes of the temperate zone release CO₂ during low tide emergence in quantities sufficient to match its metabolic production. These results indicate that CO₂ release is not hampered by the change in respiratory medium from water to air.     

Salinity and spawning of nurseryfish, <Emphasis Type="Italic">Kurtus gulliveri</Emphasis>, in the Adelaide River of northern Australia with notes on electrofishing and photos of a male carrying eggs

Nurseryfish are unique among fishes in that the males carry the fertilized eggs on a supraoccipital hook on their head. In an attempt to learn where, when, and at what salinities spawning occurs, an ichthyoplankton net was towed at 14 stations in the Adelaide River from the mouth (38 ppt) to the most upstream sections of the river (0.1 ppt). Larval nurseryfish (5–27.5 mm SL) were collected, preserved and measured, and water chemistry parameters were recorded with each sample. Larvae were found in the mid-reaches of the river during July–October, most commonly at salinities between 13.6–0.5 ppt. Salinities increased as the dry season progressed. Larvae were not taken at salinities higher than 19 ppt, nor did they occur in the upper-most reaches of the river system during this time of the dry season. What nurseryfish do and where they occur during the massive influx of freshwater runoff during the wet season (November–April) remains a mystery. Electrofishing for nurseryfish proved to be ineffective in the Adelaide River due to its extraordinary turbidity which made recovery of stunned fish difficult and fresh material of males carrying eggs remains elusive. Two historic photographs showing males carrying eggs are described.     

铜鱼和圆口铜鱼的早期发育

本文比较系统地观察了长江和汉江铜鱼和圆口铜鱼鱼卵、鱼苗的胚胎发育,记述了两种铜鱼胚胎发育的各个阶段的形态特征,比较两种铜鱼早期发育的相似性状和主要差别。这些差别主要是:雏形胚体出现的迟早、眼基和肌节出现的先后、眼睛大小、胸鳍长短以及颌须出现的迟早和长短等,依据这些差异,可以把两种铜鱼在早期发育阶段准确地区分开来。本文还对长江兴修水利枢纽与两种铜鱼资源的关系进行了调查和分析。       

长江干流和汉江的鳡鱼繁殖习性及其胚胎发育

本文报道了1961—1966年、1976—1978年和1981年在长江干流和汉江对鳡鱼的繁殖习性、产卵场的分布、产卵场的环境条件、促使产卵的外界因素和产卵规模等方面的调查研究结果。比较系统地研究了鳡鱼鱼卵、鱼苗的胚胎发育,描述了各个发育阶段的形态特征。与长江中其他产漂流性卵鱼类(特别是青鱼、草鱼、鲢和鳙)的胚胎相比较,鳡鱼具有许多明显的特点。       

Reproductive ecology of Brazilian freshwater fishes

We used metadata on nine reproductive traits of 67 species of Brazilian iteroparous, oviparous, teleost freshwater fishes to test phenotypes associations to discriminate species that only spawn in large rivers (lotic fishes) from those capable to spawn in lentic habitats (lentic fishes). We tested the hypothesis that lotic fishes present spawning migration, shorter spawning season, single spawning, no parental care, free eggs, higher relative fecundity, faster embryogenesis, and larger size, while lentic fishes present no spawning migration, longer spawning season, multiple spawning, parental care, adhesive eggs, lower relative fecundity, slower embryogenesis, and smaller size. Our analyses supported the hypothesis but not all phenotypic associations satisfied it, specifically with regard to lentic fishes or to pairs of phenotypes typical of lentic fishes. We also concluded that spawning in large rivers is a better predictor of bionomic characters than spawning in lentic habitats, and lotic fishes are specialists compared to lentic fishes.     

Location and timing of Asian carp spawning in the Lower Missouri River

We sampled for eggs of Asian carps, (bighead carp Hypophthalmichthys nobilis, silver carp H. molitrix, and grass carp Ctenopharyngodon idella) in 12 sites on the Lower Missouri River and in six tributaries from the months of May through July 2005 and May through June 2006 to examine the spatial and temporal dynamics of spawning activity. We categorized eggs into thirty developmental stages, but usually they could not be identified to species. We estimated spawning times and locations based on developmental stage, temperature dependent rate of development and water velocity. Spawning rate was higher in the daytime between 05:00 and 21:00 h than at night. Spawning was not limited to a few sites, as has been reported for the Yangtze River, where these fishes are native, but more eggs were spawned in areas of high sinuosity. We employ a sediment transport model to estimate vertical egg concentration profiles and total egg fluxes during spawning periods on the Missouri River. We did not identify substantial spawning activity within tributaries or at tributary confluences examined in this study.