The critical importance of an undammed river segment to the reproductive cycle of a migratory Neotropical fish

Biotelemetry, ichthyoplankton and genetic data can provide detailed information about the migratory dynamics and reproductive cycle of freshwater fishes. However, few studies have combined these techniques in Neotropical systems. The objective of this study was to examine the migratory and reproductive dynamics of Prochilodus costatus in the São Francisco River watershed, south‐east Brazil, by comparing the ecological importance of two rivers to the species, an undammed segment of the São Francisco River and a dammed segment of one of its main tributaries, the Pará River. In total, 215 fish were radio‐tagged over three years (2014–16). Eggs and larvae were sampled at seven locations and analysed by PCR to identify Prochilodus spp. ichthyoplankton. Most radio‐tagged individuals (97%) used the undammed segment of the São Francisco River as spawning migration route, even those captured and released in the Pará River. Fish migrated to spawn from late September to late November with the arrival of the rains and returned to feeding sites from December to May after spawning. The highest densities of fish eggs and larvae were recorded in the upper reaches of the São Francisco River during months of peak river discharge. Returning fish showed high fidelity to sites occupied before spawning migration. Fish spent roughly 71% of the year at feeding sites, 25% at spawning sites and 4% moving between them. This study provides novel information about the migratory dynamics of Neotropical fishes and underscores the key role of undammed river segments for the conservation of Neotropical migratory fish species.     

Temperature influence on the fish assemblage structure in a large lowland river, the lower Oder River, Germany

Abstract –  Water temperatures were expected to structure river fish assemblages by promoting: (i) offspring's growth and densities in warm years, (ii) spring spawning fish when temperatures increase early and (iii) improved growth and overwintering condition from elongated vegetation periods. Favourable temperature conditions in their first year should improve growth and survival in consecutive years. In the lower Oder River, Germany, fish have been sampled annually, 0+ in July since 1997, and older fish in November since 1998. This data set was analysed to test the hypotheses mentioned above. The water temperature ranged between −0.4 and 25.1 °C during the study period, resulting in substantial variations of the 29 temperature metrics derived from daily means. Cumulative degree-days until catch, in spring and during March, as well as minimum and mean temperatures in March and spring explained 63% of the variability observed. The mean total length of the most frequent 0+ fish differed significantly between years related to temperature. Increased length growth at higher temperatures was a general tendency. Total length and density of 0+ fish were significantly inversely correlated, but the potential influence of temperature on species densities was not significant. The mean fish densities varied not significantly between years. No indication was found that higher total length in the first year of life resulted in an improved growth in subsequent years.
For future research and management it has to be expected that temperature impacts become increasingly significant in regulated river systems. With decreasing hydrodynamics the significance of temperature effects will increase.     

Distribution, larval stage duration and growth of the sea lamprey ammocoetes, Petromyzon marinus L., in a highly modified river basin

Abstract  – Larval stage duration of the sea lamprey, Petromyzon marinus , in the River Mondego is estimated to last for 4 years. The number of annuli provides reliable age estimates when compared with length–frequency distributions analysis. The growth rate of the sea lamprey ammocoetes displays strong seasonal patterns, and reaches its highest value during the first 2 years of larval stage. About 69% of the length increment between hatching and metamorphosis is attained at the end of the second year. There is a longitudinal gradient associated with ammocoete distribution along the river. Relative abundance of ammocoetes decreases downstream from the Açude-Ponte dam, the first obstruction encountered by the adult sea lampreys in their upstream spawning migration along the River Mondego.     

Prespawning migratory behaviour and spawning success of sea-ranched Atlantic salmon, Salmo salar L., in the River Gudenaa, Denmark

The migratory behaviour of sea-ranched Atlantic salmon, Salmo salar L., was analysed by radio-telemetry in the River Gudenaa, Denmark. The main objectives were to: (1) estimate mortality of returning adults through the fjord; (2) observe rate of progression and migratory pattern in the fjord and river; and (3) record whether spawning occurs in the river. Forty-two returning salmon (19 males and 23 females of total body length from 60–97 cm) reared and released as smolts, were caught and equipped with external radio transmitters in the outer estuary of the River Gudenaa in 1994 and 1995. Of the tagged salmon, 18 (43%) were caught in the estuary, four (10%) were not recorded after release and 20 (47%) entered the river. The mean rate of progression through the fjord was 7.6 km d−1 (range 1.4–18.2) in 1994 and 5.4 km  d−1 (range 1.6–17.1) in 1995. Eleven salmon were alive at the onset of the spawning period. Eight were retrieved dead from the river during or after the spawning period; four with empty gonads assumed to be successful spawners, and four with intact gonads. In 1994, unsuccessful spawners (found dead with intact gonads) entered the river earlier and had a longer total migration distance in the river compared to successful spawners. This suggests that spawning success of sea-ranched salmon is associated with time of river entry and river migration length.     

Profiles of sex steroids, fecundity and spawning of a migratory characiform fish from the Paraguay–Paraná basin: a comparative study in a three-river system

This study investigated for the first time the reproductive biology of Prochilodus lineatus in a system of rivers in southeastern Brasil, relating it to the role of tributary rivers in the reproductive success of this important commercial fish in the Upper Paraná River basin, where a cascade of hydroelectric dams were deployed. Specimens were caught bimonthly in three river sites: (S1) Grande River, downstream from the Porto Colômbia dam; (S2) Pardo River; and (S3) Mogi Guaçu River. Sex steroid plasma levels, fecundity, follicular atresia, oocyte diameter and gonadosomatic index (GSI) were compared among sites. In S1, fish exhibited changes in the reproductive parameters: lower GSI, oocyte diameter and fecundity and higher follicular atresia index, when compared to S2 and S3. Frequency of maturing fish was higher in S3 and spawning was only registered in S3. In sites S2 and S3, plasma concentrations of testosterone and 17β-estradiol in females and testosterone in males showed wide variations following gonadal maturation. Fish from S1 showed few significant variations in sex steroid concentrations throughout the gonadal cycle. These results indicate that P. lineatus does not reproduce in Grande River (S1), but probably uses the Pardo River (S2) as a migratory route towards the Mogi Guaçu River (S3) where they complete gonadal maturation and spawning. Our findings contribute for understanding the reproductive biology of P. lineatus and to highlight the importance of tributaries in impounded rivers as a favourable environment for migration and spawning of fish.     

Population decline in the endemic Atlantic salmon (Salmo salar) in Kapisillit River,Greenland

Atlantic salmon, Salmo salar L., is found throughout the North Atlantic, with thousands of rivers having spawning populations. In Greenland, spawning is limited to one river in West Greenland, the Kapisillit River (64?N), and the salmon are limited to the lower few kilometres of the system. Using mark–recapture, it was estimated the parr population was 5,953 individuals, and that the population size has declined by 52% since 1959. In spite of this decline, parr density remains high, being between 0.26 and 0.62 parr/m². Using a historical age‐length key, an estimated minimum of 635 smolt will have descended to the sea in 2017. These will be caught in a fishery currently subject to no regulatory measures and fishing remains the most likely driver of the population decline. The genetically distinct population is endemic to Greenland, and managers should implement measures to conserve this genetic integrity and local biodiversity.     

Population structure and reproduction of the African bonytongue Heterotis niloticus in the Sô River-floodplain system (West Africa): implications for management

Abstract –  Population structure of the African bonytongue, Heterotis niloticus , was examined in southern Benin within Lake Hlan and a region of the Sô River floodplain located approximately 60 km downstream from the lake. Both locations support important fisheries in which Heterotis is the principal target species during the flood period. Ripe adults comprised over 40% of the population in Lake Hlan, whereas only 3.5% of individuals captured from river sites were adults. Monthly averages for the gonadosomatic index and percentages of individuals with mature gonads peaked as water levels increased during the wet season then declined during the peak flood period. Oocyte size frequency distributions from ovaries suggested a potential to produce an additional cohort in the event of nesting disruption. During the peak spawning period (May to August), between 37 and 51 active nests per hectare per month were observed in Lake Hlan. The number of larvae per nest ranged from 3953 to 6125. Lake Hlan bonytongues appear to constitute an important source subpopulation that exports new recruits to river/floodplain areas downstream where intense fisheries harvest mostly juveniles and subadults. Consequently, restriction of harvest of adult bonytongues in Lake Hlan may be essential for sustenance of commercial fishing in downstream reaches of the Sô River.     

Patterns of movement of adult northern pike (Esox lucius L.) in a regulated river

Abstract –  We performed a tagging study on one of the spawning populations of northern pike ( Esox lucius L.) of the River Kajaaninjoki in Finland. Northern pike is the main predator in many lakes and rivers in the northern hemisphere. Previous tagging studies have shown a general tendency to sedentary behaviour by pike. Whether individuals in a fish population adapt a sedentary or moving strategy may affect population persistence. To study if the spawning population consists of sedentary or moving individuals and the pattern of movements, we tagged 40 pike with radio transmitters and followed them from May 2002 to June 2003. Pike were caught and tagged during the spawning season in the mouth of the River Kajaaninjoki which flows into one of the largest lakes in Finland, Lake Oulujärvi. Our results suggest that the pike spawning population consisted of sedentary pike ( N  = 16) dwelling the whole year in the river, and moving pike ( N  = 24) which moved to Lake Oulujärvi after the spawning period. Pike exhibited homing behaviour, as most of the migrating pike returned to the same spawning area in the following year. Large pike had a higher movement rate than small pike during the summer and seasonally the movement rate was lowest during the spring and highest during the summer.     

Linking habitat characteristics with juvenile density to quantify Salmo salar and Salmo trutta smolt production in the river Sävarån,Sweden

Abstract Habitat mapping along 85 km of river was related to juvenile (15 years of electric fishing) and smolt (3 years of screw‐trapping) abundance data to estimate salmon, Salmo salar L., and sea trout, Salmo trutta L., smolt production in the River Sävarån, northern Sweden. Spawning site selection by radio‐tagged salmon (n = 12) and sea trout (n = 4) was also assessed. Fifty‐one hectares of potential spawning and nursery habitat was found in the main stem river, representing 25% of the total river area. These areas were estimated to yield 1300–7580 salmon and 630–3540 sea trout smolts based on juvenile densities, equating with 3 years of screw‐trap data (2990–5080 salmon and 680–2520 trout smolts, respectively). A hypothetical maximum production of about 19 900 salmon smolts was predicted for the river at a density of 40, 0+ salmon 100 m^?2. Tracking adults during the spawning period identified optimal and potential reproductive areas.     

Environmental DNA reveals the temporal and spatial extent of spawning migrations of European shad in a highly fragmented river basin

1. Artificial barriers on lowland rivers impede the spawning migrations of anadromous fishes, preventing access to historical spawning areas. In the cryptic European shads Alosa alosa and Alosa fallax (‘shad’ hereafter), this has resulted in population declines across their range. Conservation programmes aim to facilitate the passage of migrators over these barriers and so require baseline information on the spatial and temporal extent of current migrations.
2. Here, a shad-specific environmental DNA (eDNA) assay was used to quantify the spatial extent of shad spawning migrations in the River Severn basin, western England. This basin is characterized by the presence of multiple barriers in the lower catchment. In 2017, the eDNA assay was piloted in the River Teme, an important shad spawning tributary, and then applied in 2018 and 2019 across the lower Severn basin.
3. In all years, shad DNA was detected between mid-May and mid-June, with the maximum spatial extent of shad distribution being in early June when shad eDNA was detected upstream of weirs that were generally considered as impassable. In 2018, this included the detection of shad above the most upstream weir on the main River Severn that required individual fish to have passed six weirs.
4. Although barriers inhibit the spawning migrations of shad, this eDNA assay showed that some highly vagile individuals might be able to ascend these barriers and migrate considerable distances upstream. This suggests that efforts to increase the permeability of these barriers could result in relatively high numbers of migrating shad reaching upstream spawning areas. These results demonstrate that this eDNA assay could also be used across their range, to further quantify the spatial extent of their spawning, including in highly fragmented rivers and those where shad are believed to spawn only occasionally and are rarely observed.

     

基于年龄结构的中华鲟资源量估算方法

根据长江中的中华鲟亲鱼捕捞数据的年龄结构,推算不同年龄段中华鲟亲鱼进入长江参与繁殖的占比,首次将长江和海洋中的中华鲟同时纳入估算模型进行计算,构建了一套估算中华鲟资源量的新方法。稳态计算结果显示,在葛洲坝截流前,长江中每年有效补充量为1 882尾,长江和海洋中育龄(雌:13~34龄,雄:8~27龄)总资源量为32 260尾,其中雄鱼15 310尾,雌鱼16 950尾,每年在长江中参与繁殖的中华鲟新老股群之和(1 727尾)占总资源量的比例约5%。计算得出葛洲坝截流后长江中的中华鲟产卵繁殖容量仅为截流前的6.5%,1981年葛洲坝截留造成68%~80%的1980年老股群被阻隔在上游。结合葛洲坝截流后的捕捞数据推算了1981年后长江和海洋中的中华鲟资源量变迁过程。计算结果与捕捞数据反映的趋势一致,证明模型可靠有效。研究表明,葛洲坝截流后,随着捕捞量的减少,长江中的繁殖群体数量上升,1990年左右达到峰值(约2 200尾),随后迅速下降,2010年为170尾左右。葛洲坝截流后中华鲟产卵繁殖环境容量的大幅下降是近年来中华鲟资源量急剧下降的重要原因。     

Tributaries influence recruitment of fish in large rivers

Abstract –  Recent work demonstrates that tributary inputs are important community reorganisation points for river biota; however, no studies have examined the long-term effects of tributary inputs on fish population dynamics. This study examines nearly 40 years of young-of-year (yoy) paddlefish recruitment data to investigate the hypothesis that tributaries influence mainstem fish population dynamics. We generated hydrological variables from daily mean flow data (1965–2007) from an impounded reach of the mainstem Missouri River and from the Niobrara River, a relatively unaltered tributary, using Indicators of Hydrologic Alteration software. Three multiple regression models using natural-log transformed catch per unit effort (log cpue) as the response variable were created using (1) Missouri River-only flow variables, (2) Niobrara River-only flow variables and (3) Missouri River and Niobrara River flow variables. Flow variables from the Niobrara River explain a greater proportion of yoy paddlefish log cpue variability demonstrating that tributaries can positively impact fish population dynamics in altered rivers.     

The effects of flow on Atlantic salmon (Salmo salar) redd distribution in a UK chalk stream between 1980 and 2015

Atlantic salmon are an ecologically and economically important migratory fish in the UK, whose stocks have been declining over the past 30 years. Future climate and water use changes have the potential to alter the reproductive behaviour and distribution of salmon within a river, by restricting times and ability to access suitable spawning areas. As the survival of emergent salmon juveniles is density dependent, understanding how climate‐driven changes in flow affect the location of salmon redds is important for future conservation efforts. This study examined how flow conditions affect the distribution of redds within a UK chalk stream, the river Frome in Dorset. Sixteen years of redd distribution and flow data between 1980 and 2015 were analysed using linear mixed‐effects modelling. Generally, highest redd densities occurred within middle reaches of the main river. Mean flow during the river Frome critical migration period (October–December) did not affect the density of redds directly but affected the relationship between redd density and distance from tidal limit: redd densities were spread more uniformly throughout the river under high flow conditions, whereas redds were more aggregated in the middle river reaches under low flow conditions. Together, these findings suggest that access to upstream spawning grounds was limited under low flow conditions, which could have negative repercussions on juvenile survival. This study has revealed the distribution of redds along the river Frome for the first time and provided a basis for further study into the effects of redd distribution on subsequent juvenile life stages.     

Spawning period of a southern brown trout population in a highly unpredictable stream

Abstract –  The spawning period of brown trout ( Salmo trutta L.) was studied in the river Castril, southern Spain, by means of redd counts. This mountain stream is located near to the southern limit of the species' natural distribution range and it shows a highly unpredictable flow regime. The spawning period extended from December to mid-April and the maximum reproductive activity was in February. These results represent the latest reproduction date and the longest spawning period reported in the literature along the natural distribution range of the species. However, belated spawning in the Castril is congruent with the known latitudinal cline: the lower the latitude, the later the spawning period. Our results, along with a review of the literature on natural populations, also showed that the duration of reproduction is the longer, the lower the latitude. Spawning lasted twice as long in the main stem of the river, which is connected with a reservoir, than in the isolated reaches. These differences may be linked to the influence of the reservoir and to habitat fragmentation. We discuss and support the hypothesis that a long spawning period is an advantage for survival in unpredictable habitats. The belated and protracted spawning period found in river Castril has important implications in fisheries management. A strong research effort is needed in order to fill the critical lack of data on southern brown trout populations.     

Spatial and quantitative patterns of movement in large bull trout (Salvelinus confluentus) from a watershed in north-western British Columbia, Canada, are due to habitat selection and not differences in life history

Abstract –  Movement and habitat use by bull trout ( Salvelinus confluentus ) was examined using radio telemetry in the Morice River watershed, north-western British Columbia, between April 2000 and November 2001. Bull trout principally utilised mainstem habitat. Throughout the watershed, little movement was observed during the winter months. From June to September, movement increased and was correlated with migration into tributaries. We directly observed spawning bull trout only in tributaries of the Morice River and never in the mainstem. Many of the fish ( N  = 48) migrated to the nearest potential spawning sites, but a few fish ( N  = 7) made extensive migrations to other regions of the watershed to spawn. Fish that spawned in both years of the study returned to the same spawning location. Variation in spatial and movement data, therefore, appears to be linked to availability of suitable habitat and not differences in life history that have been observed in large bull trout.     

Reproductive ecology of two cyprinid fishes in an oligotrophic lake near the southern limits of their distribution range

Abstract –  The reproductive activities of two dominant fish species, roach Rutilus rutilus and rudd Scardinius erythrophthalmus were investigated from August 2000 to July 2002 in an oligotrophic lake, Lake Sapanca, in the northwest of Turkey. A total of 907 roach and 593 rudd individuals were examined. Macroscopic appearance of ovary, gonadosomatic index and oocyte diameter frequency distribution suggested that the spawning season of roach and rudd were from early April (13 °C) to late May (18 °C) and from early May (17.5 °C) to late June (23.5 °C), respectively. The spawning cycles of both species showed the same tendency in two successive years. The estimated mean total lengths at first maturity ( L _T50) were 12.26 and 14.98 cm for male and female roach, respectively, and 7.12 and 8.10 cm for male and female rudd, respectively. The sizes at 50% maturity corresponded to 3 years for roach and 1 year for rudd, for both sexes. The ratio of males to females was 1.3:1 and 1.2:1 for roach and rudd, respectively. The average condition factor K was 1.21 for rudd and 1.14 for roach, and it was not affected by sex or reproductive effort. Absolute fecundity varied from 7829 to 95,387 eggs between ages 3 and 9 for roach and from 1807 to 35,629 eggs between ages 1 and 7 for rudd. Comparisons of these results with earlier studies from more northern regions revealed that spawning of both species in this area started earlier, but spawning temperature and the duration of spawning was approximately the same.     

The spatial distribution of cod (Gadus morhua L.) spawning grounds in the Kattegat, eastern North Sea

Similar to many other commercial marine fish species, Atlantic cod (Gadus morhua L.) migrate towards specific sites at spawning. These temporal aggregations are generally the most targeted by the commercial fishery. The Kattegat cod has undergone a substantial reduction over the past 25 years and both stock size and spawning stock biomass have remained at very low levels since the end of the 1990s. There is, therefore, an urgent need to map and document spawning grounds still in use. In the present study, spawning sites of Atlantic cod were identified in the Kattegat using a combination of commercial and fishery independent data from 1996 to 2004. Moreover, putative spawning and non-spawning areas were also sampled before and during the reproductive season between 2002 and 2006, and the proportion of mature females together with the individual physiological status were used to validate and strengthen the spatial analyses.

The spatial analyses identified several spawning areas in the region and two areas in the southeastern part of the Kattegat which appeared to be the most important. The results showed the presence of cod spawning aggregations, although reduced in size, in areas that have been utilized for more than 25 years according to historical information. Some local spawning grounds may have also disappeared. The proportion of mature females was higher in putative spawning than in non-spawning areas (p < 0.001) and females from spawning areas had higher gonadosomatic (p < 0.05) and hepatosomatic (p < 0.001) indices than those from non-spawning areas.

Knowledge of stock spatial and temporal distribution is essential in designing recovery strategies for depleted fish populations. The unambiguous stability of the locations of spawning aggregations over time, as shown in this study, represents a useful aid in order to efficiently implement a recovery plan for the collapsing cod stock in this area.     

Multiple spawning and related variations in female reproductive parameters of amphidromous type ayu

Adult ayu Plecoglossus altivelis were sampled from the Nezugaseki River, Yamagata Prefecture to investigate the reproductive parameters and analyze their variations related to multiple spawning. Ovarian histology (mainly based on the occurrence of postovulatory follicles) indicated that female ayu can be divided into four groups, which are considered to correspond with unspawned, once spawned, twice spawned, and three (or more) times spawned individuals. Comprehensive analysis of the ovarian state with field surveys revealed that most of the female ayu achieve second spawning and that third spawning is very common in this river. Mean relative batch fecundity (RBF) for the second, third, and fourth spawning indicated 69, 42, and 22%, respectively, of the value for the first spawning, showing a decline with the number of spawning batches. Mean gonadosomatic index (GSI) values of non-ovulating fish showed a similar decline during the main spawning period (October). Condition factor (without gonads) during autumn also decreased with the number of batches spawned, suggesting that multiple spawning is achieved partly by transfer of body mass nutrients to the ovary. Size dependency in reproductive parameters such as occurrence of the multiple spawning, RBF values, and GSI values of the unspawned fish, was not evident except for GSI values in early September (generally during the early phases of ovarian development), when larger fish had larger GSI.     

Migration and life history alternatives in a high latitude species, the broad whitefish, Coregonus nasus Pallas

Abstract – We examined the expected changes in the life history trajectory of an Arctic species, broad whitefish, in the lower Mackenzie River between an anadromous population with spawning migrations of 350–450 km and a potamodromous population with spawning migrations of 5–12 km. Based on life history theory we predicted that the anadromous population would have faster growth, later age-at-maturity, and greater reproductive investment than the potamodromous population. Early in its life the anadromous population had faster growth than the potamodromous but later growth was slower. Age-at-maturity was greater in the anadromous population than the potamodromous. Reproductive investment was higher in the anadromous population. We conclude that the broad whitefish did not conclusively follow the pattern of variation observed between migratory and relatively sedentary populations in temperate zones. Fecundity was greater in the migratory population but not conclusively linked to age-at-maturity or growth-rate differences.     

Effect of habitat fragmentation on spawning migration of brown trout (Salmo trutta L.)

Abstract –  Human-induced habitat alteration is one of the main causes of the decline of freshwater fish populations. The watershed of the River Bidasoa (Spain) is an example of heavily fragmented habitat. The local brown trout ( Salmo trutta L.) population is disturbed, with evidence of poor recruitment as well as low adult densities in the main stem. Forty male and female adult migratory trout were tagged with external or internal radio transmitters and released. Fixed stations with data loggers and mobile antennae were used with daily surveys to track fish movements during the migration and spawning period (3 months). Migration distances did not exceed 10 km, and half of the fish never entered a tributary in the study area. Fragmentation because of weirs on the main stem apparently prevented fish from reaching their spawning destination. Fish that entered the tributaries were first confronted with an accessibility problem because of low discharge. However, each fish chose one tributary, without making attempts to run up in other tributaries. Once in the tributary, fish were restrained in their upstream movements by dams. The study area appeared to be isolated from the vast upper part of the watershed. Within the study area, upper parts of tributaries also seemed strongly disconnected from the main stem. This study illustrates the negative impact of river fragmentation on S. trutta migration pattern. Population sustainability can be directly affected through the low availability of spawning grounds for migratory fish. Long-term effects of fragmentation may cause reproductive isolation within watersheds, which in the case of trout also means isolated phenotypic population units.