A quantitative life history of endangered humpback chub that spawn in the Little Colorado River: variation in movement,growth, and survival

While the ecology and evolution of partial migratory systems (defined broadly to include skip spawning) have been well studied, we are only beginning to understand how partial migratory populations are responding to ongoing environmental change. Environmental change can lead to differences in the fitness of residents and migrants, which could eventually lead to changes in the frequency of the strategies in the overall population. Here, we address questions concerning the life history of the endangered Gila cypha (humpback chub) in the regulated Colorado River and the unregulated tributary and primary spawning area, the Little Colorado River. We develop eight multistate models for the population based on three movement hypotheses, in which states are defined in terms of fish size classes and river locations. We fit these models to mark–recapture data collected in 2009–2012. We compare survival and growth estimates between the Colorado River and Little Colorado River and calculate abundances for all size classes. The best model supports the hypotheses that larger adults spawn more frequently than smaller adults, that there are residents in the spawning grounds, and that juveniles move out of the Little Colorado River in large numbers during the monsoon season (July–September). Monthly survival rates for G. cypha in the Colorado River are higher than in the Little Colorado River in all size classes; however, growth is slower. While the hypothetical life histories of life‐long residents in the Little Colorado River and partial migrants spending most of its time in the Colorado River are very different, they lead to roughly similar fitness expectations when we used expected number of spawns as a proxy. However, more research is needed because our study period covers a period of years when conditions in the Colorado River for G. cypha are likely to have been better than has been typical over the last few decades.     

Parasites of native and nonnative fishes of the Little Colorado River, Grand Canyon, Arizona

A 2-yr, seasonal, parasitological study of 1,435 fish, belonging to 4 species of native fishes and 7 species of nonnative fishes from the lower Little Colorado River (LCR) and tributary creeks, Grand Canyon, Arizona, yielded 17 species of parasites. These comprised 1 myxozoan (Henneguya exilis), 2 copepods (Ergasilus arthrosis and Lernaea cyprinacea), 1 acarine (Oribatida gen. sp.), 1 piscicolid leech (Myzobdella lugubris), 4 monogeneans (Gyrodactylus hoffmani, Gyrodactylus sp., Dactylogyrus extensus, and Ligictaluridus floridanus), 4 nematodes (Contracaecum sp., Eustrongylides sp., Rhabdochona sp., and Truttaedacnitis truttae), 3 cestodes (Bothriocephalus acheilognathi, Corallobothrium fimbriatum, and Megathylacoides giganteum), and 2 trematodes (Ornithodiplostomum sp. and Posthodiplostomum sp.). Rhabdochona sp. was the only adult parasite native to the LCR. Infection intensities of Ornithodiplostomum sp. and B. acheilognathi were positively correlated with length of the humpback chub Gila cypha. Adult helminths showed a high degree of host specificity, except B. acheilognathi, which was recovered from all fish species examined but was most abundant in cyprinids. Abundance of B. acheilognathi in the humpback chub was highest in the fall and lowest in the summer in both reaches of the LCR. There was no major taxonomic difference in parasite assemblages between the 2 different reaches of the river (LC1 and LC2). Parasite community diversity was very similar in humpback chub, regardless of sampling site or time. The parasite fauna of the LCR is numerically dominated by B. acheilognathi and metacercariae of Ornithodiplostomum sp. The richest and most diverse component community occurred in a nonnative species, the channel catfish Ictalurus punctatus, but infracommunity species richness was highest in a native host, humpback chub.     

Mature males of Arctic charr,Salvelinus alpinus,release F-type prostaglandins to attract conspecific mature females and stimulate their spawning behaviour

Synopsis We examined the behavioural and biological functions of prostaglandins (PG) of F-type as a chemosensory signal during spawning in Arctic charr. Immunoassay revealed F-type PGs in water inhabited by ripe charr of both sexes performing spawning-related acts. High concentrations of immunoreactive PGFs seemed to coincide with male spawning activity. Ripe isolated males were more odorous to other charr (as assayed by EOG) than either females or immature charr. When these males received chemical cues from ovulated females, they increased their release of odorants, including F-type PGs, into the water. PGF_2α at concentrations of 0.1 nM attracted ovulated females and ripe males. It also stimulated digging behaviour in mature females. It is concluded that F-type PGs or their derivatives are released by ripe males on the spawning grounds to attract females and elicit their spawning behaviour.     

Evaluation of a conservation strategy: a spawning aggregation closure for red hind, Epinephelus guttatus, in the U.S. Virgin Islands

Many tropical reef fishes spawn in large aggregations, which are readily targeted by fishers. By the 1980s, at least two grouper spawning aggregations were eliminated by intensive fishing off the island of St. Thomas, U.S. Virgin Islands, and another aggregating species, red hind, was intensively targeted. By 1988, the average length of red hind had greatly decreased to 295 mm, and the sex ratio was extremely skewed to 15 females per male, suggesting a heavily fished stock. Since this species is a protogynous hermaphrodite, the loss of large individuals (primarily males) could potentially result in sperm limitation in spawning aggregations. In 1990, a spawning aggregation closure was implemented. By 1997, average size of red hind had increased to 395 mm and sex ratio had shifted to 4 females per male. Fish were observed aggregating only in structurally complex habitat along the insular shelf edge. This habitat type is apparently not common along most of the shelf edge off St. Thomas and may provide shelter while reducing risk of predation during aggregation periods. These data suggest that protection of spawning aggregations is a sound management strategy with considerable potential for aiding the sustainable use of reef fish resources.     

Amplitude of ecological potential: chub Leuciscus cephalus (L.) spawning in an artificial lowland canal

Chub Leuciscus cephalus (L.) is a rheophilic cyprinid which prefers lotic habitat conditions, shallow water (0.1–0.3 m) and gravel banks with moderate to high water flow (0.15–0.75m s^–1) for spawning. In contrast to these spawning requirements, a self-sustaining chub population was detected in a German lowland canal, the Oder–Havel Kanal. This canal is monotonously embanked with rip rap and almost completely lacks water flow and habitats that are suitable for riverine species. In 1999 chub spawned in the absence of water flow, in depths up to 1.28 m on allochthonous gravel with a mean diameter of 39 ± 16 mm (SD). These findings indicate a substantially higher environmental plasticity and tolerance of chub than previously reported.     

Of Travertine and Time: Otolith Chemistry and Microstructure Detect Provenance and Demography of Endangered Humpback Chub in Grand Canyon,USA

We developed a geochemical atlas of the Colorado River in Grand Canyon and in its tributary, the Little Colorado River, and used it to identify provenance and habitat use by Federally Endangered humpback chub, Gila cypha.  Carbon stable isotope ratios (δ¹³C) discriminate best between the two rivers, but fine scale analysis in otoliths requires rare, expensive instrumentation. We therefore correlated other tracers (SrSr, Ba, and Se in ratio to Ca) to δ¹³C that are easier to quantify in otoliths with other microchemical techniques. Although the Little Colorado River’s water chemistry varies with major storm events, at base flow or near base flow (conditions occurring 84% of the time in our study) its chemistry differs sufficiently from the mainstem to discriminate one from the other. Additionally, when fish egress from the natal Little Colorado River to the mainstem, they encounter cold water which causes the otolith daily growth increments to decrease in size markedly. Combining otolith growth increment analysis and microchemistry permitted estimation of size and age at first egress; size at first birthday was also estimated. Emigrants < 1 year old averaged 51.2 ± 4.4 (SE) days and 35.5 ± 3.6 mm at egress; older fish that had recruited to the population averaged 100 ± 7.8 days old and 51.0 ± 2.2 mm at egress, suggesting that larger, older emigrants recruit better. Back-calculated size at age 1 was unimodal and large (78.2 ± 3.3 mm) in Little Colorado caught fish but was bimodally distributed in Colorado mainstem caught fish (49.9 ± 3.6 and 79 ± 4.9 mm) suggesting that humpback chub can also rear in the mainstem. The study demonstrates the coupled usage of the two rivers by this fish and highlights the need to consider both rivers when making management decisions for humpback chub recovery.     

Mate choice by males of the hermit crab Pagurus filholi: Do males assess ripeness and/or fecundity of females?

Males of the hermit crab, Pagurusfilholi, often grasp the edges of shells occupied by females and drag them during the mating season. This behavior was experimentally confirmed to be a precopulatory guarding behavior displayed by males for ripe females, and males were found to recognize females which were within about 5 days of spawning. Most theoretical models for mating preference assume the choosing sex (the male in the present case) has complete reproductive information about potential mates, and predict that males will preferably choose more fecund females and/or females that will require less guarding time (i.e. that will spawn sooner) as partners. Several male-choice experiments between two ripe females, both previously guarded by other males, were carried out to examine the above predictions. Males did not prefer females of larger size, higher fecundity or with less time remaining until spawning. These results suggest that males may not have complete information about potential partners, rather that male hermit crabs may adopt a mating strategy of pairing with the first ripe female they encounter. Even with such incomplete mate assessment, males may enhance their reproductive success by recognizing ripe females that will spawn within a given time (about 5 days in the present case).     

Behavioral Tactics of Male Sockeye Salmon (Oncorhynchus nerka) under Varying Operational Sex Ratios

Previous studies have demonstrated several reproductive-behavior patterns in male salmon, including competitive and sneaking tactics, the formation of hierarchies, and non-hierarchical aggregations around ripe females. Through behavioral observations at varying spatial and temporal scales, we examined the hypothesis that operational sex ratio (OSR) determines male sockeye salmon (Oncorhynchus nerka) distribution and breeding tactics. Patterns of male distribution and behavior varied over both coarse and fine scales, associated with apparent shifts in reproductive opportunities, the physical characteristics of the breeding sites, and the deterioration of the fish as they approached death. Females spawned completely within a few days of arriving on the spawning grounds, whereas males courted the available ripe females from the date of their arrival on the spawning ground until their death. This difference in reproductive lifespans tended to elevate late-season OSRs but was partially counterbalanced by male departures and the arrival of other ripe females. The proportion of males able to dominate access to ripe females decreased and the number of large courting groups increased over the course of the season, apparently related to both increasing OSR and the deteriorating physical condition of males. However, great variation in OSR was observed within the spawning sites on a given day. OSRs were generally higher in shallow than in deep water, perhaps because larger females or more desirable breeding sites were concentrated in shallow water. The aggregations of males courting females were not stable (i.e. many arrivals and departures took place) and male aggression varied with group size. Aggression was most frequent at low OSRs and in groups of intermediate size (2–4 males per female), and much less frequent in larger groups, consistent with the needs of maximizing reproductive opportunities while minimizing unproductive energy expenditure. These results indicate that, while OSR strongly influences male distribution and behavior, other factors such as physical condition, limited movement and habitat choice may also affect male reproductive opportunities.     

Spawning Periodicity, Spawning Migration, and Size at Maturity of Green Sturgeon, Acipenser medirostris, in the Rogue River, Oregon

The Rogue River, Oregon represents one of three important spawning systems for green sturgeon, Acipenser medirostris, in North America. In this paper we describe the spawning migration, spawning periodicity, and size at maturity for green sturgeon caught in the Rogue River during 2000–2004. Green sturgeon were caught by gill net or angling; 103 individuals were tagged with radio or sonic transmitters (externally or internally). Green sturgeon caught by gill net and angling ranged from 145 cm to 225 cm total length. Histological and visual examinations of gonad tissues indicated that most green sturgeon were spawning or post-spawning adults that entered the Rogue River to spawn. Ripe individuals were caught when water temperature was 10–18°C. Specimens carrying transmitters migrated 17–105 km up river; reaches consisting of likely spawning sites were identified based on sturgeon migratory behavior. Most green sturgeon remained in the Rogue River until late fall or early winter when flows increased, after which they returned to the ocean. Eight green sturgeon (males and females) returned to the Rogue River 2–4 years after leaving, entering the river during March, April, and May when water temperatures ranged from 9°C to 16°C. None of the 103-tagged individuals entered the Rogue River during successive years. There appear to be few known natural threats to adult green sturgeon in the Rogue River. However, our data suggest that a high percentage of adults that spawn in the Rogue River (particularly males) were susceptible to harvest by commercial, Tribal, and sport fisheries after leaving the system because they were not adequately protected by maximum size limits during the period of this study. The implications of maximum size limits (or lack of size limits) to green sturgeon are discussed, and recent actions taken by Oregon and Washington Fish and Wildlife Commissions to manage green sturgeon more conservatively are presented.     

Abundance and residency dynamics of the Indo-Pacific humpback dolphin,Sousa chinensis,in the Dafengjiang River Estuary,China

Robust population size estimates are essential for informing population conservation status. Residency dynamics show population habitat use through time. Population size of Indo-Pacific humpback dolphins (Sousa chinensis) has been extensively investigated in Chinese waters, but their residency dynamics are rarely known. Mark-recapture analysis based on photo-identification records was applied to humpback dolphins in the Dafengjiang River Estuary habitat, one of the key habitats in the northern Beibu Gulf, China. Movement analyses based on lagged identification rate indicated the humpback dolphins spent, on average, 78.5 days inside and 46.9 days outside the survey area. Within the study area, the humpback dolphin abundance was 83 identifiable dolphins. A total of 353–430 humpback dolphins, estimated by POPAN modeling, were involved in this fluid habitat-use dynamic. Robust Design analysis showed strong seasonality in humpback dolphin abundance and emigration probability, implying a movement- and habitat-use pattern likely associated with spatiotemporal distribution of oceanographic characteristics and prey occurrences. Population surveys and conservation measures currently conducted in Chinese waters seldom consider seasonality in movements between habitat patches, which can be addressed by genetic analyses across habitats and cross-matching photo-identification records among neighboring habitats.     

Reproductive behaviour of a temperate serranid fish, Paralabrax clathratus(Girard), from Santa Catalina Island, California, U.S.A.

The reproductive behaviour of the kelp bass Paralabrax clathratus was studied on Santa Catalina Island, California, U.S.A. from April 2000 to September 2002. Adults formed aggregations of three to > 200 individuals, and spawning occurred within subgroups of three to 23 individuals that contained a single female. The gonado‐somatic index ( I _G) of collected ripe males (mean = 5·8%, range = 0·5–13·1%) indicated a large investment in sperm production that is common in group‐spawning fishes characterized by intense sperm competition. Spawning occurred 32 min before sunset to 120 min after sunset, and both males and females were capable of spawning multiple times during a single evening. Behavioural observations of adults and estimates of spawning periodicity from the collection of females with hydrated oocytes suggested that spawning occurred continuously throughout the summer months and showed no significant relationship with the lunar cycle. In general, the spawning behaviour of kelp bass was similar to other functionally gonochoric, group‐spawning serranids. The dynamics of P. clathratus spawning aggregations, however, were inconsistent with that of tropical reef fish spawning aggregations, including the transient spawning aggregations of some tropical serranids. Aggregation spawning appeared to be an important component of the annual reproduction of this species.     

Reproduction in an aggregating grouper,the red hind,epinephelus guttatus

Synopsis We examined the reproductive pattern of an aggregating grouper, the red hind,Epinephelus guttatus, in Puerto Rico. Macroscopic and histological examination of gonads confirmed that, although mature, ripe ovaries are found over a three-month period, spawning activity is limited to about 2 weeks each year. Females are determinate spawners and individuals are able to spawn more than once during the course of the annual spawning season. The red hind is protogynous and 50% maturity is attained at 215 mm fork length. In western Puerto Rico, spawning occurs in aggregations at several sites within loosely defined areas located towards the edge of the insular platform. Sex ratios of individuals taken by hook and line at one of the aggregation areas over a consecutive six-year period, suggest considerable intra- and inter-annual variation most likely attributable to a combination of differential ingress and egress by males and females during the course of an aggregation and to fluctuations in recruitment, differential mortality by sex and sex change among years. A comparative assessment of the reproductive patterns of seven western AtlanticEpinephelus spp. suggests that aggregation-spawning is associated with medium- to large-sized groupers, while smaller groupers do not aggregate. Mating systems vary among congeners in association with trends in male-female size dimorphism, sexual pattern and sperm competition. The short-term, localized, nature of spawning aggregations renders species with this reproductive mode particularly vulnerable to fishing pressure at spawning sites. Aggregating species, therefore, may require special management consideration.     

Competitive mate searching in male common toads, Bufo bufo

The reproductive behaviour of a population of individually marked toads Bufo bufo was studied at a pond where males outnumbered females by between four and five to one. There was intense competition between the males for mates and only 20·5 % of them bred successfully. Of the successful males, 38·5 % got mates by fighting and displacing other males from the backs of females (takeovers). Larger males enjoyed greater reproductive success because they were stronger and better able to achieve takeovers. When competing for females, some males searched at the spawn site while others searched away from spawn. The numbers searching in the two areas can be predicted by a model which assumes that unpaired males distribute themselves so that there is a spatial ESS, where individuals have equal expectations of finding a female both at and away from the spawning ground.     

Leuciscus cephalm (Linnaeus), with records of fingerling adult males, Leuciscus plemobipunctatus (Stephanidis) and their hybrids from western Greece

The spawning season of chub, Leuciscus cephalus , in western Greece starts in January-February when males are aged at least 2 + and females 3 + (rarely 2 +) years. In the R. Ermitsa (Achelos R. basin) a population is reported with fingerling adult males with developed gonads and breeding tubercles at a size of 42 72 mm s.l . Leuciscus pleurobipunctatus breeds between February and the first half of May, when males are aged at least 1 + and females 2 + (sometimes also 1 +); their life span is about 4 years. L. cephalus × L. pleurobipunctatus hybrids were found in the R. Ermitsa (five specimens) and R. Louros (one specimen). In the former the contemporary presence of adult fingerling male chub, the smallest adults of L. pleurobipunctatus , and hybrids between the two species, may be related to channel width and the human interference which has resulted in considerable disturbance of the habitat.     

Social and reproductive behavior of a captive group of walleye pollock,Theragra chalcogramma

Synopsis The social and reproductive behavior of a group of four male and seven female walleye pollock,Theragra chalcogramma, were observed in a large tank. Pollock spent most of their time swimming in a loose aggregation near the surface. Males descended from the aggregation more often than females to follow and make physical contact with other males as well as with females. The difference between males and females in the frequency of diving in our tank is consistent with the reported pattern of depth segregation of the sexes in natural pollock spawning aggregations. The frequency of social interactions increased when pollock became reproductively active and was higher at night and during twilight when most of the spawning occurred. Male interactions with females most frequently involved physical contact, while male interactions with other males were more often limited to following. There was no indication that male-male interactions result in the formation of stable social dominance relationships that determine priority of access to mates, as has been suggested previously for walleye pollock. Rather, following and contact interactions appear to promote male identification of potential mates and encounters with ripe females. The possible functional significance of male social interactions is discussed in relation to reports on natural walleye pollock spawning aggregations.     

Ovulation and Eggs of the Hagfish Eptatretus burgeri

Abstract Ripe females and males of Eptatretus burgeri were kept in cages both in the sea and in the aquaria for about 2 months of the normal spawning period around October. All the 9 ripe females in the sea ovulated all their eggs (200) and deposited 178 of them. Only 5 eggs were ovulated and deposited in the aquaria, although there were 17 ripe females and some of them were injected with various hormones in an effort to induce ovulation. No eggs were fertilized. A peculiar surface pattern of hexagonal ridges is described around the entrance of the micropylar canal on the bottom of the micropylar funnel in the ovulated eggs.     

Growth of dark chub, Zacco temmincki (Cyprinidae), with a discussion of sexual size differences

I examined the annual and seasonal growth of dark chub, Zacco temmincki, in a Japanese river. Investigation of opercular rings showed that the fish reached a maximum age of 8 years. There was no significant sexual size difference at younger ages (3–5 years), but males were larger than females at older ages (6–7 years). Annual increments of length and weight for males that were recaptured were also larger than those for females. The fish spawn from June to August. Females grew for a short period from April to May, but males on average grew for a longer period from April to August. There was no sexual difference in growth rate except during the spawning period. Annual growth rate was negatively correlated with fish length in each sex. The sexual size differences at older ages of the fish might be due to the polygynous mating system in which most mature males could not obtain females and invested for somatic growth in the spawning period, and a short growing season that was overlapped considerably with the spawning period.     

Brown trout (Salmo trutta,Linnaeus, 1758) spawning habitat improvement in the Ebro River Basin (eastern Spain)

Limestone streams in Spain are subject to calcium carbonate precipitation processes, which make the substrate very cohesive. Many populations of brown trout (Salmo trutta, Linnaeus, 1758) in those rivers show reduced production due to unsuitable spawning habitats. This study presents data on S. trutta spawning habitat improvement in the Guadalope River Basin during 2011. The main objectives of this study were to (i) modify the substrate in order to make it easier for S. trutta females to dig when spawning, (ii) test the effect of the gravel quality improvement on the habitat, (iii) analyse the effect of habitat improvement on S. trutta recruitment, and (iv) monitor habitat improvement effectiveness in the long term. Three manual measures were adopted to make the gravel suitable for S. trutta breeding: the separation of the embedded gravel using an iron bar by hitting it with a mace‐hammer (‘substrate breaking’), the spreading out of the gravel loosened by the previous action with a rake (‘raking up’), and thirdly the elimination of fine sediment depositions (‘substrate cleaning’). Results of the first year of monitoring (2012) in improved and control areas were compared. Methodology effectiveness assessment requires monitoring over a longer period, but the data from the first year after habitat improvement provide encouraging results, mainly dealing with changes in spawning habitat parameters and an increase in the S. trutta age‐0 class in the improved areas.     

Aggregation and spawning by lampreys (genus Ichthyomyzon) beneath cover

Synopsis Lampreys are generally reported to spawn in shallow water on open, gravel bottoms. During surveys in Wisconsin and Minnesota, we regularly observed aggregations of adult Ichthyomyzon c.f. gagei, I. castaneus, and I. fossor beneath such cover objects as boulders, woody debris, and, at one site, vegetation. In some cases, observations of eggs or rapid quivering by individual lampreys indicated that spawning was occurring. The literature includes scattered anecdotal reports of similar behavior in other populations of Ichthyomyzon. Our data for I. c.f. gagei suggest that aggregations beneath cover objects occur at a greater range of depths than those in the open, but that aggregations in the open can contain greater numbers of individuals. Facultative spawning beneath cover objects may permit lampreys to spawn in deep waters with swift current where spawning could not otherwise occur. Moreover, this behavior may reduce the vulnerability of spawning lampreys to some types of predators.     

Kootenai River velocities,depth, and white sturgeon spawning site selection – a mystery unraveled?

The Kootenai River white sturgeon Acipenser transmontanus population in Idaho, US and British Columbia (BC), Canada became recruitment limited shortly after Libby Dam became fully operational on the Kootenai River, Montana, USA in 1974. In the USA the species was listed under the Endangered Species Act in September of 1994. Kootenai River white sturgeon spawn within an 18‐km reach in Idaho, river kilometer (rkm) 228.0–246.0. Each autumn and spring Kootenai River white sturgeon follow a ‘short two‐step’ migration from the lower river and Kootenay Lake, BC, to staging reaches downstream of Bonners Ferry, Idaho. Initially, augmented spring flows for white sturgeon spawning were thought to be sufficient to recover the population. Spring discharge mitigation enhanced white sturgeon spawning but a series of research investigations determined that the white sturgeon were spawning over unsuitable incubation and rearing habitat (sand) and that survival of eggs and larvae was negligible. It was not known whether post‐Libby Dam management had changed the habitat or if the white sturgeon were not returning to more suitable spawning substrates farther upstream. Fisheries and hydrology researchers made a team effort to determine if the spawning habitat had been changed by Libby Dam operations. Researchers modeled and compared velocities, sediment transport, and bathymetry with post‐Libby Dam white sturgeon egg collection locations. Substrate coring studies confirmed cobbles and gravel substrates in most of the spawning locations but that they were buried under a meter or more of post‐Libby Dam sediment. Analysis suggested that Kootenai River white sturgeon spawn in areas of highest available velocity and depths over a range of flows. Regardless of the discharge, the locations of accelerating velocities and maximum depth do not change and spawning locations remain consistent. Kootenai River white sturgeon are likely spawning in the same locations as pre‐dam, but post‐Libby Dam water management has reduced velocities and shear stress, thus sediment is now covering the cobbles and gravels. Although higher discharges will likely provide more suitable spawning and rearing conditions, this would be socially and politically unacceptable because it would bring the river elevation to or in excess of 537.66 m, which is flood stage. Thus, support should be given to habitat modifications incorporated into a management plan to restore suitable habitat and ensure better survival of eggs and larvae.