RADIOTAGGING STUDIES OF BELUKHA WHALES (DELPHINAPTERUS LEUCAS) IN BRISTOL BAY, ALASKA

Research was conducted in Bristol Bay, Alaska, to determine the applicability of radiotagging to studies of behavior, distribution and movements of belukha whales. Backpack-style VHF transmitters were attached to two belukhas by pinning through the dorsal ridge. Both packages were shed after about 2 wk due to migration of the pin through the tissue. Movements of radio-tagged whales were essentially local within Kvichak Bay. Three basic respiration patterns were identified: surfacings that were grouped into breathing periods separated by longer dives; surfacings that did not occur during restricted breathing periods; and long-to-very-long surfacings separated by short-to-very-short dives. These patterns were interpreted as representing traveling, feeding and feeding or resting in very shallow water. Surface and dive interval data were used to calculate a correction factor of 2.75, which could then be applied to aerial survey counts to estimate the total number of belukha whales in the study area. Modifications to radio packages are necessary in order to increase retention time.     

GROWTH LAYER GROUPS (GLGs) IN THE TEETH OF AN ADULT BELUKHA WHALE (DELPHINAPTERUS LEUCAS) OF KNOWN AGE: EVIDENCE FOR TWO ANNUAL LAYERS

Growth layer groups (GLGs) were counted in teeth from an adult male belukha whale ( Delphinapterus leucas ) that had lived for 23 yr in captivity and was estimated to have been 14 mo at capture. As defined here, GLGs are repeating areas of alternating opaque (denser) and translucent (less dense) dentin or cementurn, or nodes at the dentin-cementum interface. A GLG involves at least one change from opaque to translucent, dense to less dense or ridge to groove, but may be further subdivided by incremental growth layers or laminations. Teeth were prepared by two techniques. Thick sections (longitudinal half sections) were examined with a dissecting microscope under reflected light. Thinner cross and longitudinal sections were x-rayed and the plates scanned with a microdensitometer. Scanning electron photomicrographs were prepared from thick sections, but SEM proved to be the least useful technique. Counts of GLGs were variable because the laminations were numerous and some surface layers had been lost from wear. Thick- and thinner-section techniques gave comparable results, and approximately 40 GLGs were counted consistently. Although captive environments are less variable than natural ones, our findings are further evidence that belukhas in the wild deposit more than one—and probably two—GLGs per year.     

CONTROL OF GERMINATION OF ALEXANDRIUM TAMARENSE (DINOPHYCEAE) CYSTS FROM THE LOWER ST. LAWRENCE ESTUARY (CANADA)

Cysts of the toxic dinoflagellate Alexandrium tamarense (Lebour) Balech 1992 from the lower St. Lawrence estuary were used in a test of the following hypotheses: (1) cyst germination is triggered by a change in temperature, and (2) germination rate varies throughout the year and is controlled by a circannual internal biological clock. Results show that cyst germination was not affected significantly by temperature of incubation over the range 1°–16° C, and light showed no significant stimulation of germination. This is supported by the lack of effect of cyst incubation conditions during evaluation of the seasonal changes in germination rate (two temperatures: 4° and 15° C, and two light conditions: darkness and 150 μmol photons·m^?2·s^?1). Thus, direct environmental control through short-term increases in temperature and exposure to light has no effect on the germination of the cysts tested. The rate of germination, observed monthly over a 16-month period, showed low germination (<20%) over most of the period tested, except for a maximum reaching more than 50% germination in August to October of the second year of the experiment. This pattern was observed for cysts both from monthly field collections and from laboratory-stored cysts kept under constant environmental conditions (4° C, in the dark). The peak in germination observed under constant environmental conditions (in the laboratory), the almost coincidental increase in cyst germination observed for the field-collected cysts, and the absence of effects of temperature and light during incubation could be explained either by a temperature-controlled cyst maturation period (the time-temperature hypothesis of Huber and Nipkow 1923) or by the presence of an internal biological clock. However, the large decline in the rate of germination 2 months after the maximum provides strong support for the biological clock hypothesis. The ca. 12-month maturation (dormancy) period observed for the laboratory-stored cysts is the longest reported for this species to our knowledge; this might be related to the low storage temperature (4° C), which is close to bottom temperatures generally encountered in this environment (0° to 6° C). Similar field and laboratory storage temperatures could explain the coincidental increase in germination rate in the fall of the second year if cyst maturation is controlled by temperature. A fraction of the laboratory-stored cysts did not follow a rhythmic pattern: A rather constant germination rate of about 20% was observed throughout the year. This continuous germination of likely mature cysts may supplement the local blooms of this toxic dinoflagellate, as these often occur earlier than peak germination observed in late summer. It seems that two cyst germination strategies are present in the St. Lawrence: continuous germination after cyst maturation, with temperature controlling the length of the maturation period, and germination controlled by a circannual internal rhythm.     

ENVIRONMENTAL FACTORS CONTROLLING ALEXANDRIUM TAMARENSE (DINOPHYCEAE) GROWTH RATE DURING A RED TIDE EVENT IN THE ST. LAWRENCE ESTUARY (CANADA)1

The dinoflagellate Alexandrium tamarense (Lebour) Balech 1985 is responsible for recurrent outbreaks of paralytic shellfish poisoning in the St. Lawrence Estuary. In July 1998, an A. tamarense red tide developed in the estuary with maximum cell concentrations reaching 2.3 × 10⁶ cells·L^?1 in brackish surface waters. To estimate the growth rate of these cells, surface water samples from different locations and days during the bloom were incubated for 5 to 9 days under in situ temperature and light conditions. Growth rates varied both spatially and temporally between 0 and 0.55 day^?1, reaching the maximum growth rate reported for this species in culture. High growth rates were measured even during the peak of the red tide, suggesting that the extremely high cell concentrations observed did not solely result from aggregation or physical concentration but also involved active cellular growth. Alexandrium tamarense cells were found over a large range of salinity (20.8–29.5 psu), but high densities and significant growth were only measured when salinity was lower than 24.5 psu. Under these conditions, the number of divisions achieved by A. tamarense was proportional to the amount of nitrate available at the beginning of the incubations, whereas variations in growth rate were apparently controlled by the availability of phosphate. We hypothesize that the ability of A. tamarense to perform vertical migrations and acquire nitrate at night pushes this species toward phosphate limitation in the St. Lawrence Estuary.     

CHARACTERISTICS OF FINBACK BALAENOPTERA PHYSALUS VOCALIZATIONS IN THE ST. LAWRENCE ESTUARY

ABSTRACT

Sounds produced by Finback Whales Balaenoptera physalus were recorded from a stationary hydrophone in the St. Lawrence Estuary from June to September. The vocalizations consisted of frequencies below 120 Hz; impulsive sounds had frequencies up to 1 kHz. Over 80% of the sounds were downsweeping calls. Frequency variations in the downsweeps were correlated with social context. Constant calls, upsweeps, wavers and a frequency and amplitude modulated call were rare and may be context specific. Vocalization rates varied with the number of animals present and context, but could not be used as a census technique. Comparisons are made with the data from other investigators in both the Northwest Atlantic and the Northeast Pacific. Frequency and time characteristics for Finback downsweeps are summarized and discussed as components important for species recognition.     

POPULATION INDEX ESTIMATES FOR THE ST. LAWRENCE BELUGAS, 1973–1995

The belugas ( Delpbinapterus leucas ) inhabiting the St. Lawrence estuary were freely hunted until 1979 and thereby much reduced in numbers. In the late 1970s concerns for their future, and for the effect of contaminants, habitat modification, and marine traffic, became acute, and they were declared "Endangered" in 1983.
The numerical progress of the population has been monitored since 1973 by continued sample surveys. Nine of 10 surveys were aerial, and of those 9, 7 used high-altitude aerial photography. Published indices of population size showed an average increase of about 17 belugas per year (SE = 4) and a 1995 smoothed value of about 650 (SE = 40) belugas. This apparent temporal trend might be open to question owing to slight variation in survey methods and in the area covered by different surveys, so factors were developed to correct for differences in survey coverage and in treatment of data and for the effect of hunting on population trajectory. Revised assessments of the population trajectory and of the present smoothed index value changed little as a result. A series of total-count surveys in 1987-1992 estimated a decline of about 6 belugas per year, and when they were combined with sample surveys flown in 1984-1995, an increase of 10 belugas per year was estimated and a 1995 standardized index of about 640 (SE = 43). It appeared that hunting in the 1970s could have consumed all of, perhaps even slightly more than, the potential for natural increase of this population.     

THE TROPHIC ROLE OF MARINE MAMMALS IN THE NORTHERN GULF OF ST. LAWRENCE

The trophic role of apex predators was evaluated in the northern Gulf of St. Lawrence ecosystem. An Ecopath model was developed for the period 1985–1987 prior to the collapse of commercially exploited demersal fish stocks in this area. Marine mammal trophic levels were estimated by the model at 4.1 for cetaceans, 4.4 for harp seals (Pagophilus groenlandicus), 4.7 for hooded seals (Cystophora cristata), 4.5 for gray seals (Halichoerus grypus), and 4.3 for harbor seals (Phoca vitulina). Harp seals were the third most important predator on vertebrate prey following large Atlantic cod (Gadus morhua) and redfish (Sebastes spp.). Different seal species preyed on different levels of the food chain. Harp seals preyed on most trophic groups, whereas larger seals, such as gray seals and hooded seals, mainly consumed higher trophic levels. The model suggested that apex predators had a negative effect on their dominant prey, the higher trophic level fish, but an indirect positive feedback on the prey of their preferred prey, mainly American plaice (Hippoglossoides platessoides), flounders, skates, and benthic invertebrates. Our results suggest that both marine mammals and fisheries had an impact on the trophic structure.     

PRESENCE OF ALGAE IN FRESHWATER ICE COVER OF FLUVIAL LAC SAINT‐PIERRE (ST. LAWRENCE RIVER,CANADA)1

Winter ice cover is a fundamental feature of north temperate aquatic systems and is associated with the least productive months of the year. Here we describe a previously unknown freshwater habitat for algal and microbial communities in the ice cover of the freshwater St. Lawrence River, Quebec, Canada. Sampling performed during winter 2005 revealed the presence of viable algal cells, such as Aulacoseira islandica (O. Müll.) Simonsen (Bacillariophyceae), and microbial assemblage growing in the ice and at the ice–water interface. Vertical channels (1–5 mm wide) containing algae were also observed. Concentrations of chl a ranged between 0.5 and 169 μg · L^?1 of melted ice, with maximal concentrations found in the lower part of the ice cores. These algae have the potential to survive when ice breakup occurs and reproduce rapidly in spring/summer conditions. Freshwater ice algae can thus contribute to in situ primary production, biodiversity, and annual carbon budget in various habitats of riverine communities.     

F(ST) and Q(ST) under neutrality

A commonly used test for natural selection has been to compare population differentiation for neutral molecular loci estimated by F_ST and for the additive genetic component of quantitative traits estimated by Q_ST. Past analytical and empirical studies have led to the conclusion that when averaged over replicate evolutionary histories, Q_ST = F_ST under neutrality. We used analytical and simulation techniques to study the impact of stochastic fluctuation among replicate outcomes of an evolutionary process, or the evolutionary variance, of Q_ST and F_ST for a neutral quantitative trait determined by n unlinked diallelic loci with additive gene action. We studied analytical models of two scenarios. In one, a pair of demes has recently been formed through subdivision of a panmictic population; in the other, a pair of demes has been evolving in allopatry for a long time. A rigorous analysis of these two models showed that in general, it is not necessarily true that mean Q_ST = F_ST (across evolutionary replicates) for a neutral, additive quantitative trait. In addition, we used finite-island model simulations to show there is a strong positive correlation between Q_ST and the difference Q_ST − F_ST because the evolutionary variance of Q_ST is much larger than that of F_ST. If traits with relatively large Q_ST values are preferentially sampled for study, the difference between Q_ST and F_ST will also be large and positive because of this correlation. Many recent studies have used tests of the null hypothesis Q_ST = F_ST to identify diversifying or uniform selection among subpopulations for quantitative traits. Our findings suggest that the distributions of Q_ST and F_ST under the null hypothesis of neutrality will depend on species-specific biology such as the number of subpopulations and the history of subpopulation divergence. In addition, the manner in which researchers select quantitative traits for study may introduce bias into the tests. As a result, researchers must be cautious before concluding that selection is occurring when Q_ST ≠ F_ST.     

DIATOM PALEOLIMNOLOGY OF TWO FLUVIAL LAKES IN THE ST. LAWRENCE RIVER: A RECONSTRUCTION OF ENVIRONMENTAL CHANGES DURING THE LAST CENTURY

Water quality degradation is a serious concern for the St. Lawrence River. While some environmental data are available for the St. Lawrence ecosystem, long-term monitoring data are generally lacking. To infer past environmental changes, we undertook a paleolimnological assessment of diatom assemblages preserved in four ²¹⁰Pb- and ¹³⁷Cs-dated sediment cores from two fluvial lakes in the river, and used diatom transfer functions to infer past shoreline habitat characteristics. At sites in Lake Saint-François, a fluvial lake downstream from Cornwall, water quality decreased this century in response to human impacts (e.g. macrophyte density and nutrient levels increased). These trends were apparent from an increase in epiphytic diatom taxa, followed by an increase in eutrophic planktonic taxa. Water quality, however, appears to have improved somewhat in response to rehabilitation measures during the last two decades. From a sediment core near Montréal (Lake Saint-Louis), we also noted a large proportion of eutrophic and epiphytic taxa, but less evidence was recorded of a recent improvement in water quality. The diatom-based inference model for habitat characteristics appeared to reconstruct environmental conditions in the St. Lawrence River during the last century. The most notable shift has been an increase in diatom taxa commonly associated with macrophyte substrates. Trends in some of the planktonic diatoms were similar to those recorded in paleolimnological investigations from Lake Ontario, but cores from the river also may be reflecting local environments. This study shows that diatom-based paleolimnological studies are possible in large river systems, if coring sites (e.g. fluvial lakes) are carefully selected.     

THE EFFECT OF VESSEL NOISE ON THE VOCAL BEHAVIOR OF BELUGAS IN THE ST. LAWRENCE RIVER ESTUARY,CANADA

During June-July 1991, we monitored the vocal behavior of belugas before, during, and after exposure to noise from a small motorboat and a ferry to determine if there were any consistent patterns in their vocal behavior when exposed to these two familiar, but different sources of potential disturbance. Vocal responses were observed in all trials and were more persistent when whales were exposed to the ferry than to the small boat. These included (1) a progressive reduction in calling rate from 3.4–10.5 calls/whale/min to 0.0 or <1.0 calls/whale/min while vessels were approaching; (2) brief increases in the emission of falling tonal calls and the theree pulsed-tone call types; (3) at distances <1 km, an increase in the repetition of specific calls, and (4) a shift in frequency bands used by vocalizing animals from a mean frequency of 3.6 kHz prior to exposure to noise to frequencies of 5.2-8.8 kHz when vessels were close to the whales.     

EPIPHYTES ON CLADOPHORA GLOMERATA IN THE GREAT LAKES AND ST. LAWRENCE SEAWAY WITH PARTICULAR REFERENCE TO THE RED ALGA CHROODACTYLON RAMOSUM (= ASTEROCYTIS SMARGDINA)1

The range of Cladophora glomerata² along the east and north shorelines of the Great Lakes and St. Lawrence Seaway extends form just east of Montreal to Thunder Bay on Lake Superior. However, it does not occur at sites sampled in Georgian Bay, the North Channel or eastern Lake Superior. The dominant epiphytes on Cladophora throughout this range are the blue-green algae, particularly Lyngbya diguetii, L. epiphytica and Chamaesiphon incrustans, which account for 53 to 90% of the cell density. The diatoms Cocconeis pediculus and Rhoicosphenia curvata contribute to much of the remaining density. The red alga Chroodactylon ramosum is a minor component of Cladophora epiphyton but is widespread in Lakes Ontario, Erie and Huron. Filament morphology, cell diameters and lengths of Chroodactylon are quite similar to those of marine forms. In addition, this alga has been reported to be quite tolerant of a wide range of salinities. Chroodactylon has been found in 33 freshwater sites throughout North America and 30 of these are from the Great Lakes or its drainage basin. Therefore, it appears possible that Chroodactylon, like Bangia atropurpurea, has originated in the Great Lakes by a migration from the Atlantic Ocean.     

BISCALITHECA KANSANA SP. N. (COENOPTERIDALES,ZYGOPTERIDACEAE), A COMPRESSION FROM THE LAWRENCE SHALE (UPPER PENNSYLVANIAN), KANSAS,U. S. A.

The organ bearing the sporangia of Biscalitheca kansana is a laminate rachis with subopposite fertile appendages, a structure previously unknown for Biscalitheca. The sporangia differ from those of the petrified Biscalitheca musata only by lacking nests of sclerotic cells. They contain spores which are called Convolutispora when found dispersed. Speculations are made that Monoscalitheca fasciculata will ultimately be regarded as a synonym of Biscalitheca musala and that some fossils called Aphlebia and Brittsia problematica may be the sterile foliage of zygopterids.