Behavior ecology of the Humboldt squid, Dosidicus gigas: insights from an animal-borne video and data logging system

Dosidicus gigas is a large pelagic cephalopod of the eastern Pacific that has recently undergone an unexpected, significant range expansion up the coast of North America. The impact that such a range expansion is expected to have on local fisheries and marine ecosystems has motivated a thorough study of this top predator, a squid whose lifestyle has been quite mysterious until recently. Unfortunately, Dosidicus spends daylight hours at depths prohibitive to making observations without significant artificial interference. Observations of this squid‟s natural behaviors have thus far been considerably limited by the bright illumination and loud noises of remotely-operated-vehicles, or else the presence of humans from boats or with SCUBA. However, recent technological innovations have allowed for observations to take place in the absence of humans, or significant human intrusion, through the use of animal-borne devices such as National Geographic‟s CRITTERCAM. Utilizing the advanced video recording and data logging technology of this device, this study seeks to characterize unknown components of Dosidicus gigas behavior at depth. Data from two successful CRITTERCAM deployments reveal an assortment of new observations concerning Dosidicus lifestyle. Tri-axial accelerometers enable a confident description of Dosidicus orientation during ascents, descents, and depth maintenance behavior - previously not possible with simple depth tags. Video documentation of intraspecific interactions between Dosidicus permits the identification of ten chromatic components, a previously undescribed basal chromatic behavior, and multiple distinct body postures. And finally, based on visualizations of spermatophore release by D. gigas and repetitive behavior patterns between squid pairs, this thesis proposes the existence of a new mating behavior in Dosidicus. This study intends to provide the first glimpse into the natural behavior of Dosidicus, establishing the groundwork for a comprehensive ethogram to be supported with data from future CRITTERCAM deployments. Cataloguing these behaviors will be useful in accounting for Dosidicus‟ current range expansion in the northeast Pacific, as well as to inform public interest in the impacts this expansion will have on local fisheries and marine ecosystems.

Social structure and underwater behavior of harbor seals in southern Monterey Bay, California

To understand harbor seal social and mating strategies, I examined site fidelity, seasonal abundance and distribution, herd integrity, and underwater behavior of individual harbor seals in southern Monterey Bay. Individual harbor seals (n = 444) were identified by natural markings and represented greater than 80% of an estimated 520 seals within this community. Year to year fidelity of individual harbor seals to southern Monterey Bay coastline was 84% (n = 388), and long-term associations (>2 yrs) among individuals were common (>40%). Consistent with these long-term associations, harbor seals were highly social underwater throughout the year. Underwater social behavior included three primary types: (1) visual and acoustic displays, such as vocalizing, surface splashing, and bubble-blowing; (2) playful or agonistic social behavior such as rolling, mounting, attending, and biting; and (3) signal gestures such as head-thrusting, fore-flipper scratch~ng, and growling. Frequency of these types of behavior was related to seal age, gender, season, and resource availability. Underwater behavior had a variety of functions, including promotion of learning and social development, reduction of aggression and preservation of social bonds by maintaining social hierarchy, and facilitation of mate selection during breeding season. Social behavior among adult males was significantly correlated with vocalization characteristics (r = 0.99, X2 = 37.7, p = 0.00087), indicating that seals may assess their competition based on underwater vocalization displays and adopt individual strategies for attracting females during breeding season based on social status. Individual mating strategies may include defending underwater territories, using scramble tactics, and developing social alliances. (PDF contains 105 pages)

Population structure, long-term connectivity, and effective size of mutton snapper (Lutjanus analis) in the Caribbean Sea and Florida Keys

Genetic structure and average long-term connectivity and effective size of mutton snapper (Lutjanus analis) sampled from offshore localities in the U.S. Caribbean and the Florida Keys were assessed by using nuclear-encoded microsatellites and a fragment of mitochondrial DNA. No significant differences in allele, genotype (microsatellites), or haplotype (mtDNA) distributions were detected; tests of selective neutrality (mtDNA) were nonsignificant after Bonferroni correction. Heuristic estimates of average long-term rate of migration (proportion of migrant individuals/generation) between geographically adjacent localities varied from 0.0033 to 0.0054, indicating that local subpopulations could respond independently of environmental perturbations. Estimates of average longterm effective population sizes varied from 341 to 1066 and differed significantly among several of the localities. These results indicate that over time larval drift and interregional adult movement may not be sufficient to maintain population sustainability across the region and that there may be different demographic stocks at some of the localities studied. The estimate of long-term effective population size at the locality offshore of St. Croix was below the minimum threshold size considered necessary to maintain the equilibrium between the loss of adaptive genetic variance from genetic drift and its replacement by mutation. Genetic variability in mutton snapper likely is maintained at the intraregional level by aggregate spawning and random mating of local populations. This feature is perhaps ironic in that aggregate spawning also renders mutton snapper especially vulnerable to overexploitation.

Age and growth of spiny dogfish (Squalus acanthias) in the Gulf of Alaska: analysis of alternative growth models

Ten growth models were fitted to age and growth data for spiny dogfish (Squalus acanthias) in the Gulf of Alaska. Previous studies of spiny dogfish growth have all fitted the t0 formulation of the von Bertalanffy model without examination of alternative models. Among the alternatives, we present a new two-phase von Bertalanffy growth model formulation with a logistically scaled k parameter and which estimates L0. A total of 1602 dogfish were aged from opportunistic collections with longline, rod and reel, set net, and trawling gear in the eastern and central Gulf of Alaska between 2004 and 2007. Ages were estimated from the median band count of three independent readings of the second dorsal spine plus the estimated number of worn bands for worn spines. Owing to a lack of small dogfish in the samples, lengths at age of small individuals were back-calculated from a subsample of 153 dogfish with unworn spines. The von Bertalanffy, two-parameter von Bertalanffy, two-phase von Bertalanffy, Gompertz, two-parameter Gompertz, and logistic models were fitted to length-at-age data for each sex separately, both with and without back-calculated lengths at age. The two-phase von Bertalanffy growth model produced the statistically best fit for both sexes of Gulf of Alaska spiny dogfish, resulting in L∞ = 87.2 and 102.5 cm and k= 0.106 and 0.058 for males and females, respectively.

Diet composition and prey selection of the introduced grouper species peacock hind (Cephalopholis argus) in Hawaii

The introduced grouper species peacock hind (Cephalopholis argus), was the dominant large-body piscivore on the Main Hawaiian Island (MHI) reefs assessed by underwater visual surveys in this study. However, published data on C. argus feeding ecology are scarce, and the role of this species in Hawaiian reef ecosystems is presently not well understood. Here we provide the first comprehensive assessment of the diet composition, prey electivity (dietary importance of prey taxa compared to their availability on reefs), and size selectivity (prey sizes in the diet compared to sizes on reefs) of this important predator in the MHI. Diet consisted 97.7% of fishes and was characterized by a wide taxonomic breadth. Surprisingly, feeding was not opportunistic, as indicated by a strongly divergent electivity for different prey fishes. In addition, whereas some families of large-body species were represented in the diet exclusively by recruit-size individuals (e.g., Aulostomidae), several families of smaller-body species were also represented by juveniles or adults (e.g., Chaetodontidae). Both the strength and mechanisms of the effects of C. argus predation are therefore likely to differ among prey families. This study provides the basis for a quantitative estimate of prey consumption by C. argus, which would further increase understanding of impacts of this species on native fishes in Hawaii.

The role of mate guarding, Male Size and Male Investment on Individual Reproductive Success in the Blue Crab, Callinectes Sapidus.

Male blue crabs, Callinectes Sapidus, guard their mates before and after mating, suggesting that the conditions regulating both types of mate guarding dictate individual reproductive success. I tested the hypothesis that large male blue crabs have advantages in sexual competition using experimental manipulations, a simulation model, and field data on crabs from mid-Chesapeake Bay between 1991-1994.

Shrimp disease management using bioactive marine secondary metabolites: an eco-friendly approach

Vibriosis caused by opportunistic and secondary bacterial pathogens is still a serious disease problem in aquaculture of the black tiger shrimp Penaeus monodon. Attempts were made for controlling shrimp bacterial disease using Marine Secondary Metabolites (MSMs). Findings indicated that the MSMs of seaweed Ulva fasciata and Dendrilla nigra are effective for controlling shrimp bacterial pathogens.

Relative importance of various predators in Clarias gariepinus fry mortality in Cameroon

To estimate the relative importance of the most common predators of Clarias gariepinus fry, increasing levels of protection were afforded to exclude amphibians, aquatic arthropods and birds. At a stocking density of 10 larvae/sq.m. in nursing ponds, fencing off amphibians resulted in a 28 per cent decrease in mortality. Holding fry in hapas to protect them from both amphibians and aquatic arthropods decreased mortality by an insignificant 5.7 per cent. Installation of bird-netting over the hapas reduced mortality by 21.7 per cent. The remaining 4.9 per cent of total mortality, which could not be explained, was attributed to opportunistic cannibalism, disease and/or handling stress. Increasing stocking density to 40/sq. m. and, thus, reducing the food available per fry increased mortality by 28.3 per cent.

Life history characteristics for silvergray rockfish (Sebastes brevispinis) in British Columbia waters and the implications for stock assessment and management

We summarize the life history characteristics of silvergray rockfish (Sebastes brevispinis) based on commercial fishery data and biological samples from British Columbia waters. Silvergray rockfish occupy bottom depths of 100−300 m near the edge of the continental shelf. Within that range, they appear to make a seasonal movement from 100−200 m in late summer to 180−280 m in late winter. Maximum observed age in the data set was 81 and 82 years for females and males, respectively. Maximum length and round weight was 73 cm and 5032 g for females and 70 cm and 3430 g for males. The peak period of mating lasted from December to February and parturition was concentrated from May to July. Both sexes are 50% mature by 9 or 10 years and 90% are mature by age 16 for females and age 13 years for males. Fecundity was estimated from one sample of 132 females and ranged from 181,000 to 1,917,000 oocytes and there was no evidence of batch spawning. Infection by the copepod parasite Sarcotaces arcticus appears to be associated with lower fecundity. Sexual maturation appears to precede recruitment to the trawl fishery; thus spawning stock biomass per recruit analysis (SSB/R) indicates that a F50% harvest target would correspond to an F of 0.072, 20% greater than M (0.06). Fishery samples may bias estimates of age at maturity but a published meta-data analysis, in conjunction with fecundity data, independently supports an early age of maturity in relation to recruitment. Although delayed recruitment to the fishery may provide more resilience to exploitation, managers may wish to forego maximizing economic yield from this species. Silvergray rockfish are a relatively minor but unavoidable part of the multiple species trawl catch. Incorrectly “testing” the resilience of one species may cause it to be the weakest member of the specie

Sex-change rules, stock dynamics, and the performance of spawning-per-recruit measures in protogynous stocks

Predicting and under-standing the dynamics of a population requires knowledge of vital rates such as survival, growth, and reproduction. However, these variables are influenced by individual behavior, and when managing exploited populations, it is now generally realized that knowledge of a species’ behavior and life history strategies is required. However, predicting and understanding a response to novel conditions—such as increased fishing-induced mortality, changes in environmental conditions, or specific management strategies—also require knowing the endogenous or exogenous cues that induce phenotypic changes and knowing whether these behaviors and life history patterns are plastic. Although a wide variety of patterns of sex change have been observed in the wild, it is not known how the specific sex-change rule and cues that induce sex change affect stock dynamics. Using an individual based model, we examined the effect of the sex-change rule on the predicted stock dynamics, the effect of mating group size, and the performance of traditional spawning-per-recruit (SPR) measures in a protogynous stock. We considered four different patterns of sex change in which the probability of sex change is determined by 1) the absolute size of the individual, 2) the relative length of individuals at the mating site, 3) the frequency of smaller individuals at the mating site, and 4) expected reproductive success. All four pat-terns of sex change have distinct stock dynamics. Although each sex-change rule leads to the prediction that the stock will be sensitive to the size-selective fishing pattern and may crash if too many reproductive size classes are fished, the performance of traditional spawning-per-recruit measures, the fishing pattern that leads to the greatest yield, and the effect of mating group size all differ distinctly for the four sex-change rules. These results indicate that the management of individual species requires knowledge of whether sex change occurs, as well as an understanding of the endogenous or exogenous cues that induce sex change.

Spatial and Temporal Distribution of Sea Turtles in the Western North Atlantic and the U.S. Gulf of Mexico from Marine Recreational Fishery Statistics Survey (MRFSS)

Systematic surveys, along with opportunistic sightings, have provided important information on sea turtle (Cheloniidae and Dermochelydae) distributions, knowledge which can help reduce the risk of harmful human interaction. In 1991 and 1992, the Marine Recreational Fishery Sta- tistics Survey (MRFSS) of the National Ma- rine Fisheries Service, NOAA, provided a unique opportunity to gain additional, synoptic information on the spatial and temporal distribution of sea turtles along the U.S. Atlantic and Gulf of Mexico coasts by asking recreational anglers if they had observed a sea turtle on their fishing trip. During the spring and summer months of those years, as water temperatures warmed, the MRFSS documented an increase in sea turtle sightings in inshore waters and in a northward direction along the U.S. Atlantic Coast and in a westward direction along the northern Gulf of Mexico. This pattern reversed in the late summer and fall months as water temperatures cooled, with sea turtles concentrating along Georgia and both coasts of Florida. Although the MRFSS did not provide species or size composition of sea turtles sighted, and effort varied depending upon location of fishing activity and time of year anglers were queried, it did provide an additional and useful means of ascertaining spatial and temporal distributions of sea turtles along these coasts.

An atlas of reproductive development in rockfishes, genus Sebastes

The genus Sebastes consists of over 100 fish species, all of which are viviparous and long-lived. Previous studies have presented schemes on the reproductive biology of a single targeted species of the genus Sebastes, but all appear to possess a similar reproductive biology as evidenced by this and other studies. This atlas stages major events during spermatogenesis, oogenesis, and embryogenesis, including atresia, in six species of Sebastes (S. alutus, S. elongatus, S. helvomaculatus, S. polyspinis, S. proriger, and S. zacentrus). Our study suggests that the male reproductive cycle of Sebastes is characterized by 11 phases of testicular development, with 10 stages of sperm development and 1 stage of spermatozoa atresia. Ovarian development was divided into 12 phases, with 10 stages of oocyte development, 1 stage of embryonic development, and 1 stage of oocyte atresia. Embryonic development up to parturition was divided into 33 stages following the research of Yamada and Kusakari (1991). Reproductive development of all six species examined followed the developmental classifications listed above which may apply to all species of Sebastes regardless of the number of broods produced annually. Multiple brooders vary in that not all ova are fertilized and progress to embryos; a proportion of ova are arrested at the pre-vitellogenic stage. Reproductive stage examples shown in this atlas use S. elongates for spermatic development, S. proriger for oocyte development, and S. alutus for embryological development, because opportunistic sampling only permitted complete analysis of each respective developmental phase for those species. The results of this study and the proposed reproductive phases complement the recommended scheme submitted by Brown-Peterson et al. (2011), who call for a standardization of terminology for describing reproductive development of fishes.

A surveillance strategy for invasive species of concern in deepwater habitats of the Northwestern Hawaiian Islands

This report describes a surveillance strategy to detect deepwater invasive species in the Northwestern Hawaiian Islands. A need for this strategy was identified in the Papahānaumokuākea Marine National Monument Management Plan and the Monument’s Draft Natural Resources Science Plan. This strategy focuses on detecting two species of concern, the octocoral Carijoa riisei and the red alga Hypnea musciformis. Most research on invasive species in the Hawaiian archipelago has focused on shallow water habitats within the limits of conventional SCUBA (0-30 m). Deeper habitats such as mesophotic reefs are much more difficult to access and consequently little is known about the distribution of deepwater invasive species or their impacts. Recent deepwater (>30 m) sightings of H. musciformis and C. riisei, in and near NWHI, respectively, have prompted a call for further research and surveillance of invasive species in deepwater habitats. This report compiles the most up to date information about these two species of concern in deepwater habitats. A literature search and conversations with subject matter experts was used to identify their current distribution, preferred habitat types, optimal detection methods and ways to efficiently sample the vast extent of NWHI. The proposed sampling strategy prioritizes survey effort where C. riisei and H. musciformis are most likely to be found. At coarse spatial scales (tens to hundreds of kilometers), opportunistic observations and distance from the Main Hawaiian Islands, a principal propagule source, are used to identify high-risk islands and banks. At fine spatial scales (meters to tens of kilometers) a habitat suitability model was developed to identify high-risk habitats. The habitat suitability model focused on habitat preferences of C. riisei, since the species is well studied and adequate data exists to map habitats. There was insufficient information to identify suitable habitat for H. muscifomis. Habitat preferences for the algae are poorly understood and there is a lack of data at relevant spatial scales to map those preferences which are known. The principal habitats identified by the habitat suitability model were ledges and the edges of rugose coral reefs, where the shade loving octocoral would likely be found. Habitat suitability maps were developed for seven atolls and banks to aid in survey site selection. The protocol relied on technical divers to conduct visual surveys of benthic habitats. It was developed to increase the efficiency of surveys, maximize the probability of detection, identify important information relevant to future surveys and standardize results. The strategy, model and protocol were tested during a field mission in 2009 at several atolls and islands in NWHI. The field mission did not detect any invasive species among deepwater habitats and much was learned to improve future surveys. Data gaps and improvements are discussed.

The effects of size-selective fisheries on the stock dynamics of and sperm limitation in sex-changing fish

Fisheries models have traditionally focused on patterns of growth, fecundity, and survival of fish. However, reproductive rates are the outcome of a variety of interconnected factors such as life-history strategies, mating patterns, population sex ratio, social interactions, and individual fecundity and fertility. Behaviorally appropriate models are necessary to understand stock dynamics and predict the success of management strategies. Protogynous sex-changing fish present a challenge for management because size-selective fisheries can drastically reduce reproductive rates. We present a general framework using an individual-based simulation model to determine the effect of life-history pattern, sperm production, mating system, and management strategy on stock dynamics. We apply this general approach to the specific question of how size-selective fisheries that remove mainly males will impact the stock dynamics of a protogynous population with fixed sex change compared to an otherwise identical dioecious population. In this dioecious population, we kept all aspects of the stock constant except for the pattern of sex determination (i.e. whether the species changes sex or is dioecious). Protogynous stocks with fixed sex change are predicted to be very sensitive to the size-selective fishing pattern. If all male size classes are fished, protogynous populations are predicted to crash even at relatively low fishing mortality. When some male size classes escape fishing, we predict that the mean population size of sex-changing stocks will decrease proportionally less than the mean population size of dioecious species experiencing the same fishing mortality. For protogynous species, spawning-per-recruit measures that ignore fertilization rates are not good indicators of the impact of fishing on the population. Decreased mating aggregation size is predicted to lead to an increased effect of sperm limitation at constant fishing mortality and effort. Marine protected areas have the potential to mitigate some effects of fishing on sperm limitation in sex-changing populations.

Examination of the foraging habits of Pacific harbor seal (Phoca vitulina richardsi) to describe their use of the Umpqua River, Oregon, and their predation on salmonids

The increase in harbor seal (Phoca vitulina richardsi) abundance, concurrent with the decrease in salmonid (Oncorhynchus spp.) and other fish stocks, raises concerns about the potential negative impact of seals on fish populations. Although harbor seals are found in rivers and estuaries, their presence is not necessarily indicative of exclusive or predominant feeding in these systems. We examined the diet of harbor seals in the Umpqua River, Oregon, during 1997 and 1998 to indirectly assess whether or not they were feeding in the river. Fish otoliths and other skeletal structures were recovered from 651 scats and used to identify seal prey. The use of all diagnostic prey structures, rather than just otoliths, increased our estimates of the number of taxa, the minimum number of individuals and percent frequency of occurrence (%FO) of prey consumed. The %FO indicated that the most common prey were pleuronectids, Pacific hake (Merluccius productus), Pacific stag-horn sculpin (Leptocottus armatus), osmerids, and shiner surfperch (Cymatogaster aggregata). The majority (76%) of prey were fish that inhabit marine waters exclusively and fish found in marine and estuarine areas (e.g. anadromous spp.) which would indicate that seals forage predominantly at sea and use the estuary for resting and opportunistic feeding. Salmonid remains were encountered in 39 samples (6%); two samples contained identifiable otoliths, which were determined to be from chi-nook salmon (O. tshawytscha). Because of the complex salmonid composition in the Umpqua River, we used molecular genetic techniques on salmonid bones retrieved from scat to discern species that were rare from those that were abundant. Of the 37 scats with salmonid bones but no otoliths, bones were identified genetically as chinook or coho (O. kisutch) salmon, or steelhead trout (O. mykiss) in 90% of the samples.