Population Status of North American Green Sturgeon, Acipenser medirostris

North American green sturgeon, Acipenser medirostris, was petitioned for listing under the Endangered Species Act (ESA). The two questions that need to be answered when considering an ESA listing are; (1) Is the entity a species under the ESA and if so (2) is the “species” in danger of extinction or likely to become an endangered species in the foreseeable future throughout all or a significant portion of its range? Green sturgeon genetic analyses showed strong differentiation between northern and southern populations, and therefore, the species was divided into Northern and Southern Distinct Population Segments (DPSs). The Northern DPS includes populations in the Rogue, Klamath-Trinity, and Eel rivers, while the Southern DPS only includes a single population in the Sacramento River. The principal risk factors for green sturgeon include loss of spawning habitat, harvest, and entrainment. The Northern DPS is not considered to be in danger of extinction or likely to become an endangered species in the foreseeable future. The loss of spawning habitat is not large enough to threaten this DPS, although the Eel River has been severely impacted by sedimentation due to poor land use practices and floods. The two main spawning populations in the Rogue and Klamath-Trinity rivers occupy separate basins reducing the potential for loss of the DPS through catastrophic events. Harvest has been substantially reduced and green sturgeon in this DPS do not face substantial entrainment loss. However there are significant concerns due to lack of information, flow and temperature issues, and habitat degradation. The Southern DPS is considered likely to become an endangered species in the foreseeable future. Green sturgeon in this DPS are concentrated into one spawning area outside of their natural habitat in the Sacramento River, making them vulnerable to catastrophic extinction. Green sturgeon spawning areas have been lost from the area above Shasta Dam on the Sacramento River and Oroville Dam on the Feather River. Entrainment of individuals into water diversion projects is an additional source of risk, and the large decline in numbers of green sturgeon entrained since 1986 causes additional concern.     

Evidence of spawning by green sturgeon, Acipenser medirostris, in the upper Sacramento River, California

This study reports the only direct evidence of spawning of green sturgeon, Acipenser medirostris, in the upper Sacramento River, CA. Two green sturgeon eggs were collected with substrate mats immediately below Red Bluff Diversion Dam. One green sturgeon larva was collected with a larval net at Bend Bridge. We concluded that green sturgeon spawn in the upper Sacramento River, both above and below RBDD. Temperature ranges in the study area (10–15°C) are similar to conditions used in successful artificial rearing of green sturgeon and do not appear to be a limiting factor to successful spawning of green sturgeon; however, suitable habitat upstream of RBDD is inaccessible when dam gates are lowered.     

Captive Breeding Programs Based on Family Groups in Polyploid Sturgeons

In species with long life cycles and discontinuous availability of individuals to reproduction, implementing a long-term captive breeding program can be difficult or impossible. In such cases, managing diversity among familiar groups instead of individuals could become a suitable approach to avoid inbreeding and increase the possibility to accomplish a breeding scheme. This is the case of several sturgeon species including the Adriatic sturgeon, whose recovery depends on the management of a few captive stocks directly descended from the same group of wild parents. In the present study, relatedness among 445 potential breeders was inferred with a novel software for pedigree reconstruction in tetraploids (“BreedingSturgeons”). This information was used to plan a breeding scheme considering familiar groups as breeding units and identifying mating priorities. A two-step strategy is proposed: a short-term breeding program, relying on the 13 remaining F0 individuals of certain wild origin; and a long-term plan based on F1 families. Simulations to evaluate the loss of alleles in the F2 generation under different pairing strategies and assess the number of individuals to breed, costs and logistical aquaculture constraints were performed. The strategy proposed is transferable to the several other tetraploid sturgeon species on the brink of extinction.     

Migration of green sturgeon, Acipenser medirostris, in the Sacramento River

Adult green sturgeon, Acipenser medirostris, were collected in San Pablo Bay, California, and surgically implanted with ultrasonic acoustic tags from 2004 to 2006. An array of automated acoustic monitors was maintained in the Sacramento River to record movements of these fish. We presumed movements to known spawning areas (based on previous green sturgeon egg collections) or areas with potential spawning habitat (characterized by substrate, flow, and temperature criteria) represented a “spawning migration.” Three separate annual “spawning migrations” were recorded involving 15 individuals. The majority of the Sacramento River migrants entered the system in the months of March and April. Two different patterns of “spawning migration” and out-migration were observed. Six individuals potentially spawned, over-summered and moved out of the river with the first fall flow event. This is believed to be the common behavior of the green sturgeon. Alternatively, nine individuals promptly moved out of the Sacramento River before 1 September, and any known flow or temperature cue. Some green sturgeon appeared to be impeded on their upstream movement by the 15 May closure of the Red Bluff Diversion Dam, and at least five passed under the dam gates during downstream migration. A delay in the closure of the Red Bluff Diversion Dam would likely allow upstream passage of spawning green sturgeon, further, the potential mortality affects of downstream passage beneath the Red Bluff Diversion Dam should be assessed. Specific protection should be also given to the large aggregation of green sturgeon located in the reach of the Sacramento River adjacent to the Glen Colusa Irrigation District pumping facility.     

Kinship affects investment by helpers in a cooperatively breeding bird

Helping behaviour in cooperative breeding systems has been attributed to kin selection, but the relative roles of direct and indirect fitness benefits in the evolution of such systems remain a matter of debate. In theory, helpers could maximize the indirect fitness benefits of cooperation by investing more in broods with whom they are more closely related, but there is little evidence for such fine-scale adjustment in helper effort among cooperative vertebrates. In this study, we used the unusual cooperative breeding system of the long-tailed tit Aegithalos caudatus to test the hypothesis that the provisioning effort of helpers was positively correlated with their kinship to broods. We first use pedigrees and microsatellite genotypes to characterize the relatedness between helpers and breeders from a 14 year field study. We used both pedigree and genetic approaches because long-tailed tits have access to pedigree information acquired through social relationships, but any fitness consequences will be determined by genetic relatedness. We then show using both pedigrees and genetic relatedness estimates that alloparental investment by helpers increases as their relatedness to the recipients of their care increases. We conclude that kin selection has played a critical role in moulding the investment decisions of helpers in this cooperatively breeding species.     

Development of new microsatellite primers for green and white sturgeon

Sixty-eight primer sets for microsatellite loci were developed from microsatellite motif enriched genomic libraries of pooled DNA from the polyploid green and white sturgeon (Acipenser medirostris and A. transmontanus). Four individuals from each species were screened for polymorphism at these loci. Forty-eight loci amplified in both species, and some exhibited species-specific amplification for white or green sturgeon (8 and 12 loci, respectively). The number of alleles per locus ranged from one to 12. At least 68% of the green and 65% of the white sturgeon loci we developed are polysomic.     

Dispersal of sibling coalitions promotes helping among immigrants in a cooperatively breeding bird

Kin selection is a major force in social evolution, but dispersal is often assumed to reduce its impact by diluting kinship. In most cooperatively breeding vertebrates, in which more than two individuals care for young, juveniles delay dispersal and become helpers in family groups. In long-tailed tits (Aegithalos caudatus), however, offspring disperse to breed and helpers are failed breeders that preferentially aid kin. Helping also occurs among immigrants, but their origins are unknown and cooperation in these cases is poorly understood. Here, we combine long-term demographic and genetic data from our study population to investigate immigration and helping in this species. We first used a novel application of parentage analysis to discriminate between immigrants and unknown philopatric recruits. We then cross-checked sibship reconstruction with pairwise relatedness estimates to show that immigrants disperse in sibling coalitions and helping among them is kin biased. These results indicate that dispersal need not preclude sociality, and dispersal of kin coalitions may help maintain kin-selected cooperation in the absence of delayed dispersal.     

Ontogenetic Behavior and Dispersal of Sacramento River White Sturgeon, <Emphasis Type="BoldItalic">Acipenser Transmontanus</Emphasis>, with a Note on Body Color

Synopsis We studied Sacramento River white sturgeon, Acipenser transmontanus, in the laboratory to develop a conceptual model of ontogenetic behavior and provide insight into probable behavior of wild sturgeon. After hatching, free embryos initiated a low intensity, brief downstream dispersal during which fish swam near the bottom and were photonegative. The weak, short dispersal style and behavior of white sturgeon free embryos contrasts greatly with the intense, long dispersal style and behavior (photopositive and swimming far above the bottom) of dispersing free embryos of other sturgeon species. If spawned eggs are concentrated within a few kilometers downstream of a spawning site, the adaptive significance of the free embryo dispersal is likely to move fish away from the egg deposition site to avoid predation and reduce fish density prior to feeding. Larvae foraged on the open bottom, swam <1 m above the bottom, aggregated, but did not disperse. Early juveniles initiated a strong dispersal with fish strongly vigorously swimming downstream. Duration of the juvenile dispersal is unknown, but the strong swimming likely disperses fish many kilometers. Recruitment failure in white sturgeon populations may be a mis-match between the innate fish dispersal and post-dispersal rearing habitat, which is now highly altered by damming and reservoirs. Sacramento River white sturgeon has a two-step downstream dispersal by the free embryo and juvenile life intervals. Diel activity of all life intervals peaked at night, whether fish were dispersing or foraging. Nocturnal behavior is likely a response to predation, which occurs during both activities. An intense black-tail body color was present on foraging larvae, but was weak or absent on the two life intervals that disperse. Black-tail color may be an adaptation for avoiding predation, signaling among aggregated larvae, or both, but not for dispersal.     

Comparative stable isotope analyses of green sturgeon (Acipenser medirostris) and white sturgeon (A. transmontanus) in the San Francisco estuary

Green sturgeon (Acipenser medirostris) and white sturgeon (A. transmontanus) are closely related, sympatric species that inhabit the San Francisco estuary. Green sturgeon have a more marine life history but both species spawn in the Sacramento River and reside for some duration in San Francisco Bay. These sturgeons are of conservation concern, yet little is known about their dietary competition when they overlap in space and time. To examine evidence of dietary differentiation, we collected whole blood and blood plasma from 26 green sturgeon and 35 white sturgeon in San Francisco Bay. Using carbon and nitrogen stable isotope analyses, we compared their relative trophic levels and foraging locations along the freshwater to marine gradient. Sampling blood plasma and whole blood allowed comparison of dietary integration over shorter and longer time scales, respectively. Plasma and whole blood δ¹³C values confirmed green sturgeon had more marine dietary sources than white sturgeon. Plasma δ¹⁵N values revealed white sturgeon fed at lower trophic levels than green sturgeon recently, however, whole blood δ¹⁵N values demonstrated the two species fed at the same trophic level over longer time scales. Larger individuals of both species had higher δ¹³C values than smaller individuals, reflecting more marine food sources in adulthood. Length did not affect δ¹⁵N values of either species. Isotope analyses supported the more marine life history of green than white sturgeon and potentially highlight a temporary trophic differentiation of diet between species during and preceding the overlapping life stage in San Francisco Bay.     

Partitioning kin groups of broodstock in the critically endangered Chinese sturgeon,Acipenser sinensis Gray 1835

Pedigrees of broodstock with unknown relationship of the critically endangered Chinese sturgeon, Acipenser sinensis, was evaluated using microsatellite markers to facilitate genetic management in restocking programs with small broodstock size. We characterized the distributions of relatedness values to reconstruct kin groups in four hatchery families with known pedigrees using microsatellites. The distributions of relatedness values for kin classes were used for partitioning full sibling groups of wild A. sinensis broodstock kept in two hatcheries, resulted in 13 full sibling clusters, four of which containing 62% of all the wild individuals. This indicates high probability of choosing close related breeder pairs in random mating, thus selective breeding is necessary to minimize inbreeding and maintain genetic diversity. This study provides a useful tool for genetic management in conservation programs of A. sinensis in aim of preserving self‐sustained wild populations.     

Multiple paternity and kin recognition mechanisms in a guppy population

Help directed toward kin (nepotism) is an important example of social behaviour. Such helping behaviour requires a mechanism to distinguish kin from nonkin. The prevailing kin recognition hypothesis is that when familiarity is a reliable cue of relatedness, other mechanisms of recognition will not evolve. However, when familiarity is an unreliable cue of relatedness, kin recognition by phenotype matching is instead predicted to evolve. Here we use genetic markers to show that guppies (Poecilia reticulata) from a population in a tributary of the Paria River in Trinidad are characterized by a high degree of multiple mating with 95% of broods having more than one sire and some dams having offspring sired by six males. These levels of multiple mating are the highest reported among live-bearing fishes. The mean relatedness of brood-mates was 0.36 (as compared to 0.5 for full-siblings). Therefore, familiarity does not seem to be a reliable mechanism to assess full-sibling relatedness. Using two-choice behavioural trials, we found that juveniles from this population use both phenotype matching and familiarity to distinguish kin from nonkin. However, we did not find strong evidence that the guppies use these mechanisms to form shoals of related individuals as adults, which is similar to results from other guppy populations in Trinidad. The use of both familiarity and phenotype matching is discussed in the context of the Paria River guppy population's mating system and ecology. Overall, these data provide support for the kin recognition hypothesis and increase our understanding of the evolution of kin recognition systems.     

Atlantic and shortnose sturgeons of the Hudson River: common and divergent life history attributes

The Hudson River estuary supports substantial number of Atlantic sturgeon, Acipenser oxyrinchus, and shortnose sturgeon, Acipenser brevirostrum. Both species have complex life cycles that have been studied sporadically in the past 50 years. The life cycle of the shortnose sturgeon may be divided into four life intervals: non-spawning adults, spawning adults, eggs and larvae, and juveniles. The life cycle of the Atlantic sturgeon is reviewed in six intervals: non-spawning adults, female spawners, male spawners, eggs and larvae, early juveniles, and late juveniles. Both species are long-lived, mature at advanced age, have rapid and similar growth during the first few years of life, feed on generally similar taxa, use deep channel habitats for all life intervals, and have complex migratory patterns with distinct, seasonal, concentration areas. Atlantic and shortnose sturgeons differ, however, in ages and sizes at maturity, maximum size, timing and location of spawning, migratory behaviors, and management. Use of marine habitats and long-distance coastal migrations are restricted to Atlantic sturgeon, but some evidence indicates that large Atlantic sturgeon juveniles reside in riverine habitats along the Atlantic coast during warm months. Movements and habitat use by both sturgeons in the Hudson River estuary contrasts with the spatial segregation of the species reported in other river systems. Juvenile shortnose sturgeon and early juvenile Atlantic sturgeon have almost the same distributions in the Hudson River estuary during all seasons. During this period of co-occurrence, both species are very similar in size, grow at about the same rate, feed on similar foods, and share deep, channel habitats. Adult shortnose sturgeon distribution overlaps with the distribution of juvenile Atlantic sturgeon, and the latter commence river emigration at a size comparable to co-occurring adult shortnose sturgeon. Life history information on the Hudson River sturgeons substantiates the need to carefully conserve these species because of vulnerability to exploitation and habitat disruption.     

Behavior, movements, and habitat use of adult green sturgeon, Acipenser medirostris, in the upper Sacramento River

We conducted the first continuous shipboard tracking of southern Distinct Population Segment green sturgeon, Acipenser medirostris, in the Sacramento River. Tracking of adult green sturgeon occurred between river kilometer (rkm) 434.8 and 511.6, a section of the putative spawning grounds located near Red Bluff, California. The recorded positions of acoustically tagged green sturgeon were analyzed using First Passage Time analysis to determine differences in habitat use between suitable and non-suitable habitats. Classification and Regression Tree modeling was used to determine explanatory inputs attributable to above average habitat use. Green sturgeon exhibited above average habitat use at five sites, identified as potential spawning aggregate sites. Three types of movements (holding, milling, and directed) could be categorized from tracks. Lastly, we show that green sturgeon while on the spawning grounds exhibit a high degree of mobility throughout the spawning grounds, often making large movements between specific habitat units. Our study illustrates how the application of shipboard tracking can be useful for describing movement, behavior and habitat utilization at a spatial scale not achieved by stationary acoustic monitors.     

Ontogenetic Behavior and Migration of Gulf of Mexico Sturgeon, Acipenser oxyrinchus desotoi, with Notes on Body Color and Development

We observed Suwannee River Gulf sturgeon, Acipenser oxyrinchus desotoi, in the laboratory and found free embryos (first interval after hatching) hid under rocks and did not migrate. Thus, wild embryos should be at the spawning area. Larvae (first interval feeding exogenously) initiated a slow downstream migration, and some juveniles (interval with adult features) continued to migrate slowly for at least 5 months, e.g., a 1-step long larva-juvenile migration. No other population of sturgeon yet studied has this migration style. A conceptual model using this result suggests wild year-0 sturgeon have a variable downstream migration style with short-duration (short distance) migrants and long-duration (long distance) migrants. This migration style should widely disperse wild fish. The model is supported by field studies that found year-0 juveniles are widely dispersed in fresh water to river km 10. Thus, laboratory and field data agree that the entire freshwater reach of river downstream of spawning is nursery habitat. Foraging position of larvae and early juveniles was mostly on the bottom, but fish also spent hours holding position in the water column, an unusual feeding location for sturgeons. The holding position of fish above the bottom suggests benthic forage in the river is scarce and fish have evolved drift feeding. The unusual migration and foraging styles may be adaptations to rear in a river at the southern limit of the species range with poor rearing habitat (low abundance of benthic forage and high summer water temperatures). Suwannee River Gulf sturgeon and Hudson River Atlantic sturgeon, A. o. oxyrinchus, are similar for initiation of migration, early habitat preference, and diel migration. The two subspecies differ greatly for migration and foraging styles, which is likely related to major differences in the quality of rearing habitat. The differences between Atlantic sturgeon populations show the need for geographical studies to represent the behavior of an entire species.     

Reach specific use of spawning habitat by adult green sturgeon (Acipenser medirostris) under different operation schedules at Red Bluff Diversion Dam

Mature green sturgeon, Acipenser medirostris, enter rivers along the western coast of North America in late winter to late spring and migrate upriver to spawning sites. After spawning, they may leave the river or spend the summer and autumn holding in deep pools before departing from the river with the onset of winter rains. Evidence exists that the seasonal Red Bluff Diversion Dam (RBDD) was an obstacle to the upriver migration of green sturgeon in the Sacramento River in Central California. We compared the migratory movements of green sturgeon under three different dam operation schedules, including post‐decommissioning, to assess the impact of this management action. The proportion of green sturgeon carrying acoustic transmitters that moved above the RBDD was higher when the gates were closed on June 15, one month later than the historical closure date of May 15, and increased again after the dam was decommissioned. The application of statistical analyses (generalized linear and additive mixed models) to the detection records of green sturgeon highlighted an improvement in connectivity after dam decommissioning. The data also indicate that interannual variation in river condition is an important driver of sturgeon presence on the spawning grounds.     

Cryptic sociality in rattlesnakes (Crotalus horridus) detected by kinship analysis

Research on social behaviour has largely concentrated on birds and mammals in visually active, cooperatively breeding groups (although such systems are relatively rare) and focused much less on species that rarely interact other than for mating and parental care. We used microsatellite markers to characterize relatedness among aggregations of timber rattlesnakes (Crotalus horridus), a putatively solitary reptile that relies heavily on chemical cues, and found that juveniles and pregnant females preferentially aggregate with kin under certain conditions. The ability to recognize kin and enhance indirect fitness thus might be far more widespread than implied by studies of animals whose behaviour is primarily visually and/or acoustically mediated, and we predict that molecular markers will reveal many additional examples of 'cryptic' sociality.     

Offspring-driven local dispersal in female sand lizards (Lacerta agilis)

We report on a field study in which determinants of female breeding dispersal (i.e. the shift in the mean home range coordinates between successive breeding events) was investigated. Offspring were released in full sib groups (or half sib ones if there was within-clutch multiple paternity) at a separation distance from the females that varied between 'families'. This allowed for analysis of 'offspring nearness' effects on maternal dispersal. When a female's offspring were released more closely to her, she responded with greater dispersal. Furthermore, when the data set was truncated at 100 m maternal-offspring separation distance at offspring release (because perception at longer distances is likely to be unrealistic), maternal dispersal resulted in greater separation distance between female and offspring in the following year. A corresponding analysis for juveniles revealed no effect of maternal nearness on offspring dispersal but identified a significant effect of clutch size, to our surprise with dispersal declining with increasing clutch size. We discuss this result in a context of the 'public information hypothesis' (reinterpreted for juveniles in a nonsocial foraging species), suggesting that conspecific abundance perhaps acts as an indicator of local habitat quality. Thus, our analysis suggests a microgeographic structuring of the adult female population driven by genetic factors, either through inbreeding avoidance, or from simply avoiding individuals with a similar genotype regardless of their pedigree relatedness, while a nongenetic factor seems more important in their offspring.     

Routes to indirect fitness in cooperatively breeding vertebrates: kin discrimination and limited dispersal

Hamilton demonstrated that the evolution of cooperative behaviour is favoured by high relatedness, which can arise through kin discrimination or limited dispersal (population viscosity). These two processes are likely to operate with limited overlap: kin discrimination is beneficial when variation in relatedness is higher, whereas limited dispersal results in less variable and higher average relatedness, reducing selection for kin discrimination. However, most empirical work on eukaryotes has focused on kin discrimination. To address this bias, we analysed how kin discrimination and limited dispersal interact to shape helping behaviour across cooperatively breeding vertebrates. We show that kin discrimination is greater in species where the: (i) average relatedness in groups is lower and more variable; (ii) effect of helpers on breeders reproductive success is greater; and (iii) probability of helping was measured, rather than the amount of help provided. There was also an interaction between these effects with the correlation between the benefits of helping and kin discrimination being stronger in species with higher variance in relatedness. Overall, our results suggest that kin discrimination provides a route to indirect benefits when relatedness is too variable within groups to favour indiscriminate cooperation.     

Unscreened Water-Diversion Pipes Pose an Entrainment Risk to the Threatened Green Sturgeon,Acipenser medirostris

Over 3,300 unscreened agricultural water diversion pipes line the levees and riverbanks of the Sacramento River (California) watershed, where the threatened Southern Distinct Population Segment of green sturgeon, Acipenser medirostris, spawn. The number of sturgeon drawn into (entrained) and killed by these pipes is greatly unknown. We examined avoidance behaviors and entrainment susceptibility of juvenile green sturgeon (35±0.6 cm mean fork length) to entrainment in a large (>500-kl) outdoor flume with a 0.46-m-diameter water-diversion pipe. Fish entrainment was generally high (range: 26–61%), likely due to a lack of avoidance behavior prior to entering inescapable inflow conditions. We estimated that up to 52% of green sturgeon could be entrained after passing within 1.5 m of an active water-diversion pipe three times. These data suggest that green sturgeon are vulnerable to unscreened water-diversion pipes, and that additional research is needed to determine the potential impacts of entrainment mortality on declining sturgeon populations. Data under various hydraulic conditions also suggest that entrainment-related mortality could be decreased by extracting water at lower diversion rates over longer periods of time, balancing agricultural needs with green sturgeon conservation.