Reproductive Ecology and Host Utilization of Four Sympatric Bitterling (Acheilognathinae,Cyprinidae) in a Lowland Reach of the Harai River in Mie,Japan

Bitterling are fishes that use freshwater mussels for oviposition. The reproductive ecology and pattern of mussel utilization of four sympatric species of bitterling, Acheilognathus rhombeus, A. tabira tabira, Tanakia lanceolata, and T. limbata, were investigated in a lowland river with seven sympatric mussel species. Three bitterling species are spring spawners with overlapping spawning seasons. A. rhombeus is an autumn-spawning species and is temporally isolated in its reproduction from the other species. Ovipositor length during oviposition of T. limbata and T. lanceolata was short, while those of A. tabira tabira and A. rhombeus were long. Most T. limbata inhabited near-shore areas, whereas the two other spring-spawning species were distributed across the entire river. All bitterling species used Inversidens brandti, Obovalis omiensis and Inversiunio jokohamensis as spawning hosts, but not the other mussel species available. T. lanceolata, A. tabira tabira and A. rhombeus showed spawning preferences for O. omiensis and I. brandti. However, T. limbata did not show clear preferences for any of the mussel species they used. A. t. tabira showed a significant preference for large I. brandti in offshore areas, while the other spring-spawning bitterling showed a preference for mussels inshore. These results are discussed in the context of reproductive resource partitioning.     

The reproductive ecology of two subspecies of the bitterling Rhodeus atremius (Cyprinidae, Acheilognathinae)

The reproductive ecology of two endangered subspecies of the bitterling Rhodeus atremius was investigated in two rivers in Okayama and Fukuoka Prefectures, Japan. Several other bitterling species, R. ocellatus, Tanakia limbata, T. lanceolata, Acheilognathus tabira and A. rhombeus, are also found in these rivers. Spawning of both R. atremius subspecies was recorded between late March and August 2007, peaking between April and July. Females of R. atremius began to mature at small body size (minimum 25 mm). They had small clutch sizes (maximum 16 eggs) and developed a short ovipositor (mean 13.6 mm), which was used to deposit a bulb-shaped egg of comparatively large size (3.2 mm³, major and minor axes 2.8 and 1.5 mm). Clutch size, ovipositor length and egg size appear to be a function of female body size. Both the body size and ovipositor length of R. atremius with ripe ova and the size and shape of the eggs varied seasonally. Rhodeus a. suigensis achieved maturity at a smaller body size, had a longer ovipositor and produced smaller eggs, with the minor axis being wider relative to the major axis than those of R. a. atremius. Rhodeus atremius may be adapted for depositing small numbers of eggs in a much shallower position inside the gills of smaller host mussels than other bitterling species.     

Predicting distributions of seven bitterling fishes in northern Kyushu,Japan

The distributions of seven bitterling species and subspecies—Tanakia lanceolata, T. limbata, Acheilognathus tabira nakamurae, A. rhombeus, Rhodeus ocellatus kurumeus, R. ocellatus ocellatus, and R. atremius atremius—in northern Kyushu were predicted using generalized linear models (GLMs) in order to provide information helpful for conserving native bitterlings and preventing the expansion of alien bitterling species. Predictions were made according to the following procedure: (1) a set of GLMs for each species was formulated using environmental data from 710 sites that were derived using digital maps and GIS software, from which the best fit model for each species was selected using the Akaike information criterion for predicting the fish occurrence, (2) model performance was evaluated based on the receiver-operating characteristics (ROC) analysis using occurrence and environmental data from 362 sites, and (3) potential distributions of the bitterling were analyzed using the best fit models and environmental data for 1,272 sites, of which 200 data points without occurrence data were prepared. The best fit models revealed that 4–6 environmental factors were important in predicting seven bitterling distributions, which was supported by the area under the ROC curve (AUC) values of these fishes ranging from 0.753 to 0.927. The AUC values in model evaluation were significantly greater than 0.5 for six fishes, suggesting the moderate accuracies of these best fit models for predicting the fish distributions. These predictive models can be used for evaluating potential native bitterling richness and the potential distribution expansion of an alien subspecies.     

Bayesian inference supports the host selection hypothesis in explaining adaptive host specificity by European bitterling

Generalist parasites have the capacity to infect multiple hosts. The temporal pattern of host specificity by generalist parasites is rarely studied, but is critical to understanding what variables underpin infection and thereby the impact of parasites on host species and the way they impose selection on hosts. Here, the temporal dynamics of infection of four species of freshwater mussel by European bitterling fish (Rhodeus amarus) was investigated over three spawning seasons. Bitterling lay their eggs in the gills of freshwater mussels, which suffer reduced growth, oxygen stress, gill damage and elevated mortality as a result of parasitism. The temporal pattern of infection of mussels by European bitterling in multiple populations was examined. Using a Bernoulli Generalized Additive Mixed Model with Bayesian inference it was demonstrated that one mussel species, Unio pictorum, was exploited over the entire bitterling spawning season. As the season progressed, bitterling showed a preference for other mussel species, which were inferior hosts. Temporal changes in host use reflected elevated density-dependent mortality in preferred hosts that were already infected. Plasticity in host specificity by bitterling conformed with the predictions of the host selection hypothesis. The relationship between bitterling and their host mussels differs qualitatively from that of avian brood parasites.     

Genetic diversification of the Kanehira bitterling Acheilognathus rhombeus inferred from mitochondrial DNA,with comments on the phylogenetic relationship with its sister species Acheilognathus barbatulus

The Kanehira bitterling, Acheilognathus rhombeus, is a freshwater fish, discontinuously distributed in western Japan and the Korean Peninsula. Unusually among bitterling it is an autumn-spawning species and shows developmental diapause. Consequently, the characterization of its evolutionary history is significant not only in the context of the fish assemblage of East Asia, but also for understanding life-history evolution. This study aimed to investigate the phylogeography of A. rhombeus and its sister species Acheilognathus barbatulus, distributed in China, using a mitochondrial analysis of the ND1 gene from 311 samples collected from 50 localities in Japan and continental Asia. Phylogenetic analysis revealed that A. barbatulus is included in A. rhombeus and genetically closer to Japanese A. rhombeus than to Korean A. rhombeus. Divergence of Korean A. rhombeus and A. barbatulus from Japanese A. rhombeus was estimated to be from the late Pliocene (3.44 Mya) and the early Pleistocene (1.98 Mya), respectively. Each event closely coincided with the time of the Japan Sea opening. Japanese A. rhombeus comprised seven lineages: three in Honshu and four in Kyushu. One lineage in central Kyushu was genetically closer to the Honshu lineages than to other lineages in northern Kyushu. Divergence of Japanese lineages was estimated to be from the early to middle Pleistocene (0.55–0.93 Mya), during a period of geological and paleoclimatic change, including volcanic activity. Population expansion in the late Pleistocene (<0.10 Ma) was suggested in many of the lineages, which accords with other freshwater fishes. Biogeographically the ancestral A. rhombeus/A. barbatulus was likely to have repeatedly colonized Japan from the continent through land bridges in the late Pliocene and the early Pleistocene. However, the close genetic relationship between Japanese A. rhombeus and A. barbatulus suggests another possibility, with the second colonization occurring in reverse, from Japan to China. The small genetic distance between them indicates that the colonization occurred later than colonization events of other freshwater fishes, including other bitterling species.     

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.     

Avoidance of host resistance in the oviposition-site preferences of rose bitterling

A contemporary outcome of dynamic host–parasite coevolution can be driven by the adaptation of a parasite to exploit its hosts at the population and species levels (parasite specialisation) or by local host adaptations leading to greater host resistance to sympatric parasite populations (host resistance). We tested the predominance of these two scenarios using cross-infection experiments with two geographically distant populations of the rose bitterling, Rhodeus ocellatus, a fish brood parasite of freshwater mussels, and four populations of their mussel hosts (two Anodonta woodiana and two Unio douglasiae populations) with varying degrees of geographic sympatry and local coexistence. Our data support predictions for host resistance at the species level but no effect of local coexistence between specific populations. Rhodeus ocellatus showed a preference for allopatric host populations, irrespective of host species. Host mussel response, in terms of ejection of R. ocellatus eggs, was stronger in the more widespread and abundant host species (A. woodiana) and this response tended to be higher in sympatric populations. These outcomes provide support for the importance of host resistance in bitterling oviposition-site decisions, demonstrating that host choice by R. ocellatus is adaptive by minimizing egg ejections. These findings imply that R. ocellatus, and potentially other bitterling species, may benefit from exploiting novel hosts, which may not possess appropriate adaptive responses to parasitism.     

Females solicit sneakers to improve fertilization success in the bitterling fish (Rhodeus sericeus)

In most species there is greater variance in reproductive fitness among males than females. A consequence is the evolution of alternative male mating tactics, including sneaking behaviour whereby males attempt to mate with females courted by other males, which could undermine female choice. In bitterling (Rhodeus sericeus), fish that lay their eggs on the gills of living freshwater mussels, sneaking is common and successful. Here we show that female bitterling can significantly improve their fertility with an increase in the number of spawning partners. Females perform conspicuous behaviours associated with spawning more frequently close to sneakers, and spawn more eggs close to high-quality sneakers. This is the first time that females have been shown to engage in behaviour that increases the probability of sneaking, and raises the possibility of a sexual conflict over the number and source of ejaculates during spawning in this species.     

Hypoxia tolerance of three native and three alien species of bitterling inhabiting Lake Kasumigaura,Japan

Differences in hypoxia tolerance among three native and three alien bitterling species were examined by means of field surveys and aquarium experiments. I caught fish in minnow traps and measured environmental factors such as dissolved oxygen and current velocity once a month between June and November 2006 at 30 points around Lake Kasumigaura, Japan. In addition, aquarium experiments were carried out to compare aquatic surface respiration thresholds among the bitterling species. Much more individuals of an alien species, Rhodeus ocellatus ocellatus, was caught at the least oxygen range than expected in the field, and showed the highest hypoxia tolerance in the laboratory experiments. Another alien bitterling, Acheilognathus rhombeus, also showed higher hypoxia tolerance than the three native bitterling species. Lake Kasumigaura is well known for eutrophication and water masses with low dissolved oxygen have often been observed there. Differences in hypoxia tolerance among bitterling species might have affected bitterling assemblage structure in Lake Kasumigaura, which is currently characterized by the dominance of R. o. ocellatus.     

The genetic status of two subspecies of Rhodeus atremius, an endangered bitterling in Japan

Rhodeus atremius is an endemic bitterling fish from Japan comprising two endangered subspecies, R. a. atremius and R. a. suigensis. The latter subspecies, which had dramatically declined in last decades, was reported to contain two distinct mtDNA lineages. In order to estimate the optimized units for conservation management, the genetic structure of R. atremius was inferred by analyzing mtDNA and microsatellite markers (MS). Allelic richness and heterozygosity of MS in R. a. suigensis was less than half that in R. a. atremius. In R. a. suigensis, not only within-population genetic diversity but also among-population genetic divergence was low, with the exception of population Ah1, while the diversity was high and the divergence showed isolation by distance in R. a. atremius. In mtDNA and MS, R. a. suigensis concordantly formed a single lineage, while R. a. atremius encompassed four mtDNA lineages, two of which were completely admixed into one group on the basis of MS. In population Ah1 a striking introgression between the two subspecies was suggested by a Bayesian-based assignment test, with the presence of mtDNA haplotype of R. a. atremius. Contrary to the prevailing theory, R. a. suigensis corresponds to a single conservation unit, while three units seem appropriate for R. a. atremius. In addition, low genetic diversity of R. a. suigensis might have arisen mainly as a result of recent bottlenecks before population fragmentation, followed by current anthropogenic effects. Genetic introgression in population Ah1 was probably the result of human transplantation of R. a. atremius.     

Pheromone-Induced Spawning of Pacific Herring

A spawning pheromone in the milt (semen) and testes of the Pacific herring,Clupea harengus pallasi,is thought to facilitate school spawning of this species. We found that responsiveness to the spawning pheromone was variable among ripe fish (milt-producing or ovulated). Measurement of five principle reproductive steroids in the free form and five steroids in conjugated forms in the plasma of male fish early in the spawning season (newly ripe fish) showed that elevated plasma levels of 3α,17α-dihydroxy-5β-pregnan-20-one and 17α-hydroxyprogesterone coincided with responsiveness to the spawning pheromone in these fish; levels of other steroids did not differ. In contrast, responsiveness to the pheromone by female fish later in the spawning season (ripe-and-holding fish) coincided with lower levels of glucuronated 17α,20β-dihydroxyprogesterone and a lower gonadosomatic index. We suggest that these differences indicate a more advanced mature reproductive state in the responsive individuals among both the newly ripe male and the ripe-and-holding female fish. We found no differences in the level of cortisol in the blood of the herring that could be correlated with differences in pheromonal responsiveness. We conclude that differences in responsiveness to the spawning pheromone coincide to some extent with levels of reproductive maturation but probably not with recent stress.     

Reproductive ecology of the striped bitterling Acheilognathus cyanostigma (Cyprinidae: Acheilognathinae)

The reproductive ecology of the striped bitterling Acheilognathus cyanostigma was investigated in a small pond in Mie Prefecture, central Honshu, Japan. A. cyanostigma was the only species of bitterling in the study pond, where only a single species of mussel Anodonta woodiana was present. Spawning of A. cyanostigma was recorded between early April and early July, peaking between late April and mid-June in 2003. Ovipositor length during the spawning period ranged from 19.2 to 42.8 mm, and was positively correlated with female body length, but did not show significant seasonal variation. The eggs are elliptical with a volume of approximately 1.5 mm³. Egg size correlates positively with female body size, and both egg shape and volume changed significantly with season. The embryos were located on host mussel gills approximately 30 mm from the exhalant siphon and were found more frequently on the inner rather than the outer mussel gills. The possible ecological significance of these observations is discussed in the context of the adaptations of A. cyanostigma for utilizing mussels for oviposition.     

A taxonomic revision of the Antennaria rosea (Asteraceae: Inuleae: Gnaphaliinae) polyploid complex

TheAntennaria rosea species complex is circumscribed to contain four subspecific taxa. A complete synonymy and key to the subspecies ofA. rosea is presented. The following new combinations are proposed:Antennaria rosea subsp.arida (E. Nels.) R. Bayer,A. rosea subsp. confinis (Greene) R. Bayer, andA. rosea subsp. pulvinata (Greene) R. Bayer. The affinities ofA. rosea to other species ofAntennaria are discussed and a key to separateA. rosea from related polyploid complexes is provided.     

Reproductive characteristics of the ocean leatherjacket, Nelusetta ayraudi

The ocean leatherjacket (Nelusetta ayraudi) is one of the largest members of the family Monacanthidae. Distributed throughout inshore waters around the southern half of Australia, this schooling species supports substantial commercial and recreational fisheries. N. ayraudi do not conform to either of the general reproductive modes reported within the family, but exhibit characteristics of both strong social reproductive behaviour and of being promiscuous and polygynous. Sexual dimorphism, with males and females exhibiting differing colouration and body shapes, and ripe ovaries being an order of magnitude larger than ripe testes, are characteristic of social reproductive behaviour and pair spawning. In contrast, high batch fecundity (mean of 320 oocytes per gram of body weight), similar sizes and ages at sexual maturity (350 mm and 2.5 years respectively) and the formation of large spawning aggregations in offshore waters are characteristic of being promiscuous and polygynous. Similar to many other coastal marine species off the east and west coasts of Australia, N. ayraudi are partial spawners during the austral winter months with spawning restricted to the part of their distribution that is towards the upper area of the prevailing currents. It is hypothesized that N. ayraudi off eastern Australia have evolved a life-history strategy whereby fish move northwards through time, spawning occurs in these more northern waters and the southerly flowing Eastern Australian Current facilitates dispersal of eggs and larvae southwards. The reproductive characteristics described provide various options to fishery managers who wish to enhance the sustainability of the fishery through increased egg production. These include spatial and temporal fishing closures to protect breeding fish during the spawning period, the protection of juveniles through either inshore area closures, improving the selectivity of fishing gears and/or regulated minimum legal lengths.     

Ecology of some species of catfishSynodontis (Pisces: Mochocidae) in the Kpong Headpond in Ghana

Synopsis Some aspects of the ecology of the freshwater catfishSynodontis (Pisces: Mochocidae) in Kpong Headpond (Ghana) were studied. Five species were encountered, namelySynodontis schall, S. gambiensis, S. ocellifer, S. velifer andS. eupterus. S. ocellifer andS. velifer were not recorded in the area before impoundment and might have entered the headpond as eggs or young from the main Volta Lake. Length: weight relationships for the two most important species can be described by the equations: Log W = - 1.76 + 3.12log L (maleS. schall), Log W = -1.33 + 2.89log L (femaleS. schall) and Log W = - 0.39 + 2.07log L (S. gambiensis, sexes combined).Synodontis is omnivorous and generally browses on benthic deposits. Adults feed mostly on chironomids, plant material and aquatic insects. The genus displays sexual dimorphism by having a short urino-genital papilla in most adult males. ForS. schall, one season of major spawning activity occurs per year from mid to late September with first time spawners between 20.0–30.0 cm SL. TheSynodontis species studied lay between 2000 to 209000 eggs per spawning season.     

The reproductive ecology of the southern red tabira bitterling Acheilognathus tabira jordani in Japan

The life history, population and reproductive variables of the southern red tabira bitterling Acheilognathus tabira jordani were investigated in a lowland reach of the River Ohara in Shimane Prefecture, western Honshu, Japan. Acheilognathus t. jordani , like all other species of bitterling, lays its eggs on the gills of freshwater mussels. It was the only species of bitterling present in the study reach, and three species of bivalve mussel were available to it for spawning: Anemina arcaeformis, Anodonta lauta and Corbicula leana . Spawning by A.t. jordani was recorded between early April and early July in 2003 and began at a size of 38· 0 mm standard length ( L _S) in the 1+ age class. Ovipositor length ( L _OP) during oviposition was positively correlated with female L _S, and showed significant seasonal variation, with a mean ± s.d. L _OP of 27· 5 ± 5· 3 mm ranging from 16· 8 to 42· 0 mm during the spawning period, which was shorter than that of a previously studied A. t. tabira population. Eggs of this subspecies are relatively long and elliptic in shape, with a volume of c. 2· 4 mm³. Egg number correlated positively with female L _S and both egg shape and volume changed significantly with season. The population size of adults was estimated to be 850 individuals, and comprised age 0+ to 3+ individuals with L _S ranging from 12· 0 to 72· 2 mm. The population sex ratio was significantly female biased, with seven females: three males. Egg shape and size and L _OP during oviposition in the present A. tabira population may be the result of local adaptations to the mussel species utilized and no competition with other bitterling species for spawning sites.     

A phylogenetic groundplan of the specific complex Dactylis glomerata

A phytochemical examination of the specific complex of Dactylis glomerata L. based on flavonoid patterns, has led to a proposed evolutionary scheme. Two phyla are pointed out, originating from diploid ancestors related respectively to D.g. subsp, aschersoniana and D.g. subsp. smithii. The temperate component of the specific complex is monophyletic, derivating from a D.g. subsp, aschersoniana-like diploid ancestor. On the other hand, chemical data support a polyphyletic origin for the geographically mediterranean part of the complex with several subphyla originating from D.g. subsp. aschersoniana-like, smithii-like and secondarily juncinella-like diploids.