VARIABILITY IN THE DIET OF HARBOR PORPOISES (PHOCOENA PHOCOENA) IN SCOTTISH WATERS 1992–2003

Interannual, seasonal, and regional variation in the diet of porpoises, Phocoena phocoena , in Scottish (UK) waters was studied using stomach contents of animals stranded between 1992 and 2003. Most samples came from the east coast (including many porpoises killed by bottlenose dolphins), with smaller numbers from the west coast and from Shetland. The most important prey types, in terms of contribution by number and mass, were whiting ( Merlangius merlangus ) and sand eels (Ammodytidae). Multivariate analysis confirmed the existence of regional, seasonal, and interannual variation in diet, as well as differences ( i. e. , biases) related to cause of death. These differences were further explored using univariate analyses. Sand eels were more important in the summer months (quarters 2 and 3) and on the east coast, whereas gadids were more important in winter and in the Shetland area. Some, but not all, observed trends in the numerical importance and size of prey taken were consistent with trends in abundance and size of fish taken during research trawl surveys. There was some evidence that porpoises <1 yr old took more gobies (Gobiidae) and shrimps than did older porpoises. Clupeids (herring Clupea harengus and sprat Sprattus sprattus ) formed a relatively small proportion of the diet, but their importance varied from year to year. Although possible methodological biases prevent firm conclusions, it appears that the importance of clupeids in porpoise diet may have decreased since the 1960s, mirroring the decline in North Sea herring abundance. The recovery of the North Sea herring stock in recent years is not as yet reflected in porpoise diet.     

LIFE IN THE FAST LANE: THE LIFE HISTORY OF HARBOR PORPOISES FROM THE GULF OF MAINE

We describe the life history of harbor porpoises in the Gulf of Maine by examining 239 animals killed in gill net fisheries and comparing these findings with the results of previous studies from the Bay of Fundy. Most female porpoises matured at age three and became pregnant each year thereafter. Reproduction was strictly seasonal, with ovulation, conception, and parturition occurring in the spring and early summer. The oldest specimen in the sample was 17 yr of age, but most individuals were younger than 12. The findings are similar to those of earlier studies from the Bay of Fundy and support the hypothesis that these animals form a single population. Harbor porpoises represent one end of a continuum of odontocete life histories that spans a wide diversity of strategies. In comparison with other, large odontocetes, harbor porpoises mature at an earlier age, reproduce more frequently, and live for shorter periods.     

DIEL VARIATION IN ECHOLOCATION BEHAVIOR OF WILD HARBOR PORPOISES

The echolocation rate and behavior of wild harbor porpoises were studied using a harbor porpoise click detector (POD) deployed close to the bottom at 40 m depth in Scottish waters, UK, April—June 2001. Echolocation variables were compared among four diel phases; morning, day, evening, and night. The echolocation encounter rate, the minimum interclick interval per train, and the proportion of echolocation click trains with a minimum interclick interval below 10 msec were all significantly higher at night than during the day. The variation in echolocation rate implies that porpoises increased their echolocation rate and/or visited the depth of the POD more often at night than during the day. Further, the changes in minimum interclick interval per train suggest that they used their echolocation for foraging or investigating objects at a close range to a higher extent, and acoustically explored the environment at greater distances at night than during the day.     

DIVING BEHAVIOR AND PERFORMANCE OF HARBOR PORPOISES, PHOCOENA PHOCOENA, IN FUNKA BAY, HOKKAIDO, JAPAN

In order to monitor the diving behavior of free-ranging cetaceans, microdataloggers, with pre-programmed release mechanisms, were attached to the dorsal fins of two female harbor porpoises ( Phocoena phocoena ) in Funka Bay, Hokkaido, Japan, in 1994. The two loggers were successfully recovered and a total of 141 h of diving data (depth and water temperature in 4,671 dives) was obtained. Both porpoises dived almost continuously, rarely exhibiting long-term rest at the surface. Maximum dive depths were 98.6 m and 70.8 m, respectively, with more than 70% of diving time at 20 m or less. Most shallow dives were V-shaped with no bottom time. The V-shaped dives were significantly shallower in dive depth and shorter in dive duration than U-shaped dives. Descent rate was not constant during a dive. The deeper the dive depths, the faster the mean descent and initial descent rates. This suggests that porpoises have anticipated the depth to which they will dive before initiating the dive itself.     

BLOOD CHEMISTRY OF WILD HARBOR PORPOISES PHOCOENA PHOCOENA (L.)

Blood chemistry values were measured from 31 harbor porpoises ( Phocoena phocoena ) released from herring weirs in the Bay of Fundy, Canada. Sodium, phosphorus, calcium, chloride, magnesium, total protein, albumin, globulin, urea, cholesterol, serum osmolality, and alanine aminotransferase levels fell within ranges reported for captive harbor porpoises and other odontocetes. Glucose, potassium, creatine kinase, aspartate aminotransferase, hemoglobin, thyroxine, bilirubin, and alkaline phosphatase levels were generally higher than those reported for captive odontocetes. Cortisol and creatinine levels were significantly higher ( P = 0.0356 and 0.0174, respectively) in porpoises that were handled for longer periods and fitted with electronic tags (mean cortisol value = 314 ± 107 nmol/L, mean creatinine value = 94 ± 14 μmol/L) than those receiving roto-tags (mean cortisol value = 222 ± 82 nmol/L, mean creatinine value = 75 ± 15 μmol/L). A more controlled study of duration of handling, time spent out of water, degree of manipulation, and method of tag attachment is required to identify which factors are responsible for the stress response.     

HEMATOLOGY VALUES OF WILD HARBOR PORPOISES (PHOCOENA PHOCOENA) FROM THE BAY OF FUNDY,CANADA

Clinical hematology values were determined for 29 harbor porpoises (Phocoena phocoena) released from herring weirs in the Bay of Fundy, Canada. Erythrocyte values exhibited narrow ranges, but there was a high degree of individual variability in counts of white blood cells. Total white cell counts ranged from 2.6 to 15.5 ± 10⁹/liter, with an overall mean of 6.5 ± 2.7 ± 10±/liter. There were significant differences among reproductive classes in mean values of total red blood cells, hematocrit (HCT), mean cell volume (MCV), mean cell hemoglobin (MCHC) and total monocyte count. Wild harbor porpoises had fewer white blood cells, lower relative total number of neutrophils and lymphocytes, and higher percentages of monocytes and eosinophils than reported in the literature for captive porpoises. Compared to published values for other odontocetes, the hemograms of harbor porpoises were most similar to those of Pacific white-sided dolphins (Lagenorhynchus acutus). These hematology data represent a baseline from free-ranging harbor porpoises that can be used as a reference for long-term monitoring of the health of this population and as a tool for rehabilitation facilities.     

EFFECT OF THE SOUND GENERATED BY AN ACOUSTIC HARASSMENT DEVICE ON THE RELATIVE ABUNDANCE AND DISTRIBUTION OF HARBOR PORPOISES (PHOCOENA PHOCOENA) IN RETREAT PASSAGE, BRITISH COLUMBIA

We describe an experiment conducted to assess the impact of the sound generated by an acoustic harassment device (AHD) on the relative abundance and distribution of harbor porpoises ( Phocoena phocoena ) in Retreat Passage, British Columbia. During control periods when the AHD was inactive, the mean number of porpoises observed in the study area was 0.39 for broad area scans conducted with the naked eye and 0.48 for narrow sector scans conducted with binoculars. Abundance declined precipitously when the AHD was activated, to 0.007 porpoises per broad area scan and 0.018 per narrow sector scan. The mean number of porpoise resightings while tracking their movements also declined from 12.2 to 13.6 per sighting during control periods to 1.1–1.9 per sighting when the AHD was activated, which suggested that the few porpoises that ventured into the study area spent less time within it when the AHD was activated. The effect of the AHD diminished with distance. No porpoises were observed within 200 m of the AHD when it was activated. The number of sightings and resightings observed when it was activated was less than 0.2% of the number expected had there been no AHD effect at a range of 200–399 m, 1.4% the number expected at a range of 400–599 m, varied between 2.5% and 3.3% of the number expected at a range of 600–2,499 m, and was 8.1% the number expected at a range of 2,500–3,500 m, which suggested that the impact of the AHD extended beyond our maximum sighting range of 3.5 km.     

ECHOLOCATION RATES OF TWO HARBOR PORPOISES (PHOCOENA PHOCOENA)

The rate of occurrence of click trains of two harbor porpoises ( Phocoena phocoena ) were counted during 14 nights. We developed an echolocation signal detection system that was harnessed to a porpoise and activated a light when the animal emitted an echolocation signal. This device, referred to as a click-light, detects echolocation signals above 150 dB re 1 μPa in the 28–180 kHz range. Echolocation rates, i.e., occurrences of click trains, changed frequently, ranging from 0 to 25 per minute. Echolocation rates were affected by feeding, individual difference, and enclosure type such as the net enclosure and the pool. The porpoise echolocation rates seemed to show acclimation.     

BILATERAL DIRECTIONAL ASYMMETRY OF THE APPENDICULAR SKELETON OF THE HARBOR PORPOISE (PHOCOENA PHOCOENA)

Directional asymmetry ( DA ) of the lengths, diameters, and masses of the scapula, humerus, radius, and ulna were analyzed on a sample of 213 harbor porpoises from Denmark and West Greenland. The levels of DA were consistent across yearlings and older animals, mature and immature animals, sexes, and populations. All investigated variables showed significant DA favoring the right side. For the pooled sample, DA of lengths ranged from 0.2% SE 0.1 (ulna) to 1.2% SE 0.1 (scapula). DA s of diameters were 1.5% SE 0.2 for the humerus, 0.6% SE 0.1 for the radius, and 1.3% SE 0.2 for the ulna. DA of mass ranged from 2.8% SE 0.5 (humerus) to 4.3% SE 0.7 (ulna). The humerus and ulna had significantly larger mean diameter/ length ratios on the right side than the left, making them more robust. The large DA of scapula length indicates larger muscle mass associated with the right flipper, while the generally more robust right humerii and ulnae may be designed for higher levels of mechanical stress. These DA s and the examples of lateralized behavior recorded in cetaceans, point to the existence of lateralized use of the flippers at the population level in harbor porpoises and possibly other cetacean species.     

GEOGRAPHICAL VARIATION IN METRIC SKULL CHARACTERS AMONG PROPOSED SUBPOPULATIONS AND STOCKS OF HARBOR PORPOISE, PHOCOENA PHOCOENA, IN THE WESTERN NORTH ATLANTIC

A total of 433 skulls of harbor porpoises, Phocoena phocoena , were examined to evaluate geographical variation within the western North Atlantic, using analysis of covariance and discriminant analysis based on 45 measurements of skull components. The relationship between condylobasal length and body length was sigmoid in both sexes. Condylobasal length is generally less in adult males than in adult females.
Despite some overlap, morphological differences between different regions and even adjacent local stocks could be recognized by ANCOVA and discriminant analysis. The percentage of correct classification by discriminant functions was 65.2% for the four regional samples: (1) North Canadian Atlantic (which includes Gulf of St. Lawrence and Newfoundland), (2) Bay of Fundy, (3) Gulf of Maine-New York, and (4) Maryland-South Carolina. It was 65.9% for the four local geographical areas (10s of km) within the Bay of Fundy; Quoddy-region porpoises revealed a closer relationship with the southwestern Nova Scotia population than with other adjacent stocks. The results at this level imply that gene flow is restricted to some degree even among closely adjacent geographical units. Management of this species should aim, therefore, for conservation of a variety of local stocks.     

REACTIONS OF CAPTIVE HARBOR PORPOISES (PHOCOENA PHOCOENA) TO PINGER-LIKE SOUNDS

Pingers on gill nets can reduce bycatch of harbor porpoises. If harbor porpoises habituate to pingers, the effect may be reduced or lost. Two captive harbor porpoises were exposed to three sound types. All sounds were in the frequency band from 100 kHz to 140 kHz, 200 ms long, and presented once per 4 s. The source level was 153 dB re 1 μPa RMS at 1 m. Each session consisted of a 10‐min presound, a 5‐min sound, and a 10‐min postsound period. Behavior was recorded on video and on dataloggers placed on the dorsal fin of one animal. The loggers recorded heart rate, swimming speed, dive duration, and depth. The animals responded most strongly to the initial presentations of a sound. Surface time decreased, the heart rate dropped below the normal bradycardia, and echolocation activity decreased. The reactions of both animals diminished rapidly in the following sessions. Should the waning of responsiveness apply to wild animals, porpoises may adapt to the sounds but still avoid nets, or the bycatch may increase after some time. The success of long‐term use of pingers may then depend on the variety of sounds and rates of exposure.     

EFFECTS OF MEAL SIZE ON OTOLITH RECOVERY FROM FECAL SAMPLES OF GRAY AND HARBOR SEAL PUPS

Recovered otoliths from pinniped feces provide valuable information on diet composition and prey size. We studied the effect of meal size on otolith recovery from the feces of one harbor and eight gray seal pups. Each of 11 experiments comprised a half-ration meal, a period of fecal collection, a 1.5-or double-ration meal again followed by a period of fecal collection. A significantly lower percentage of Atlantic herring otoliths were recovered from half-ration meals (25%± 12.5% in the harbor seal, 8.6%± 6.9% in eight gray seals) than from 1.5- or double-ration meals (62.5%± 3.1 % in the harbor seal, 32.8%± 23.5% in gray seals). Meal size also significantly affected the percentage of Atlantic cod otoliths recovered from gray seal feces (65.0%± 26.3% from half ration, 98.3%± 2.9% from 1.5 ration). For both size meals, recovered cod otoliths were more significantly eroded than herring otoliths. The development of correction factors to account for the effects of digestion will need to consider the distribution of meal sizes of free-ranging pinnipeds.     

THE DISTRIBUTION, ABUNDANCE AND SELECTED PREY OF THE HARBOR SEAL, PHOCA VITULINA CONCOLOR, IN SOUTHERN NEW ENGLAND

The seasonal distribution and abundance of harbor seals occurring south of Maine were documented by counting the number of seals at traditional haulout locations. The average number of seals counted during each survey in Massachusetts and New Hampshire was 3,560 ± 255 (95% CI), 1983–1987. The maximum number of seals counted on any individual survey was 4,736 individuals. Fifty percent of all the surveys since January 1985 have resulted in counts greater than 4,000 seals reflecting a 27% increase in the abundance of seals in our study area since that date. Seventy-five percent of the seals in southern New England are located at haulout sites on Cape Cod and Nantucket Island. The largest aggregation of seals in the eastern United States occurs mid-winter at Monomoy Island and adjacent shoals. A single high count of 1,672 seals occurred at this site during the study period. An additional 271–374 seals were also counted in Rhode Island, Connecticut and eastern Long Island Sound during surveys conducted in March 1986 and 1987. The American sandlance Ammodytes americanus was the single dominant prey item of harbor seals in waters adjacent to Cape Cod based on the modified frequency of occurrence of each prey species in scat samples collected from three haulout sites on Cape Cod between 1984–1987. During January and February sandlance was the near exclusive prey item at Monomoy (99%, n= 80). During March and April, the frequency of Atlantic herring Clupea harengus increased in the scat samples at this site. Regional differences in the diet of seals reflect distinct prey communities throughout the study area. Since 1986, the percent occurrence and importance of sandlance in the diet of seals has decreased, reflecting an overall decrease in abundance of this prey species in waters adjacent to Cape Cod. In spite of fluctuations in abundance, and regional differences in the diet of seals throughout the study area, sandlance still comprised a minimum 55% of the total prey species of harbor seals throughout the study area.     

INTRASPECIFIC VARIATION IN SEX ALLOCATION IN A SIMULTANEOUS HERMAPHRODITE: THE EFFECT OF INDIVIDUAL SIZE

Within a population of simultaneous hermaphrodites, individuals may vary in both their current reproductive investment (biomass invested in gonads) and in how they allocate that investment between male and female function. In the chalk bass, Serranus tortugarum, estimates of both reproductive allocation and reproductive success as a male and a female can be made for individuals of different sizes. As individuals increase in size, their investment in gamete production increases, and there is a shift in allocation to a stronger female bias. Spawning frequency as a female in pair spawnings and as a male in both pair spawning and streaking (an alternative mating tactic) does not vary with individual size. As a result, larger individuals should release more sperm or eggs per spawn. Size-assortative pair spawning in this species leads to larger individuals having higher potential returns in total male reproductive success than smaller individuals, which should lead to increases in absolute levels of sperm production in larger individuals when individuals compete for fertilizations through sperm competition. However, smaller individuals contribute a smaller proportion of the sperm released in spawns with multiple spawners and thus are under more intense sperm competition than larger individuals, which should select for increases in male allocation in smaller individuals, all else equal. A local-mate-competition (LMC) model predicts that these factors select for increasing absolute male and female investment with individual size but a relative shift to more female-biased allocation as individual size increases. These predictions are supported by gonadal data. The predictions of average male allocation from the quantitative LMC model were 21.6% and 25.7%, whereas the collections averaged 21.3%. This close agreement of both the mean male allocation and its relative shift with individual size between model and data support the hypothesis that size-specific shifts in sex allocation in this species represent an adaptive response to patterns of mating success and sperm competition.     

DELETERIOUS MUTATIONS AND GENETIC VARIATION FOR FLOWER SIZE IN MIMULUS GUTTATUS

Abstract.— Mimulus guttatus is a wildflower that exhibits substantial genetic variation in flower size. Here, we test the hypothesis that this variation is caused by deleterious mutations maintained through mutation‐selection balance. The deleterious‐mutation model predicts that rare, partially recessive alleles will be the primary source of variation. We test this prediction by measuring the change in the mean flower size (ΔM) and the directional dominance of flower size (ΔB) within a selection experiment. If variation is due to rare (partially) recessive alleles, ΔB/ΔM is expected to be positive and exceed one. However, we obtain negative values for ΔB/ΔM from three independent selection lines. This result is statistically inconsistent with the deleterious‐mutation model.     

鄱阳湖及其支流长江江豚种群数量及分布

在1997 年11 月至1998 年11 月的134 d 中, 大致按一年四季, 分4 次对鄱阳湖及其主要支流中的长江江豚种群数量、数量的季节变动、分布、行为、江豚栖息地环境、人类活动对江豚的影响进行了考察。江豚的分布主要集中在鄱阳湖湖区, 赣江、信江、抚河等主要支流的中下游和支流入湖的湖口附近。冬、春、秋3 个季节鄱阳湖水系长江江豚种群数量的估计值分别为91 头、431 头和260头。即, 鄱阳湖江豚的种群数量约为100～400 头,其种群数量随季节、水位、鱼类资源的变化而呈现出相应的变化。     

内蒙古草原蝗虫个体大小及生活型划分的探讨

根据内蒙古锡林河流域草原蝗虫的形态测量指标,对２３种蝗虫进行了个体大小及生活型量化划分的探讨,结果表明,２３种蝗虫可以划分为大、中、小３个类群,生活型表现为从典型的地栖型到典型的植栖型的渐为特征。个体在小与生活型紧密相关,个体大的种类常为地本型,而个体小的种类多为植栖型。     

THE BETA-BINOMIAL MODEL: ACCOUNTING FOR INTER-TRIAL VARIATION IN REPLICATED DIFFERENCE AND PREFERENCE TESTS

Binomial tests are commonly used in sensory difference and preference testing under the assumptions that choices are independent and choice probabilities do not vary from trial to trial. This paper addresses violations of the latter assumption (often referred to as overdispersion) and accounts for variation in inter-trial choice probabilities following the Beta distribution. Such variation could arise as a result of differences in test substrate from trial to trial, differences in sensory acuity among subjects or the existence of latent preference segments. In fact, it is likely that overdispersion occurs ubiquitously in product testing. Using the Binomial model for data in which there is inter-trial variation may lead to seriously misleading conclusions from a sensory difference or preference test. A simulation study in this paper based on product testing experience showed that when using a Binomial model for overdispersed Binomial data, Type I error may be 0.44 for a Binomial test specification corresponding to a level of 0.05. Underestimation of Type I error using the Binomial model may seriously undermine legal claims of product superiority in situations where overdispersion occurs. The Beta-Binomial (BB) model, an extension of the Binomial distribution, was developed to fit overdispersed Binomial data. Procedures for estimating and testing the parameters as well as testing for goodness of fit are discussed. Procedures for determining sample size and for calculating estimate precision and test power based on the BB model are given. Numerical examples and simulation results are also given in the paper. The BB model should improve the validity of sensory difference and preference testing.     

AN INTERACTION BETWEEN ENVIRONMENTAL TEMPERATURE AND GENETIC VARIATION FOR BODY SIZE FOR THE FITNESS OF ADULT FEMALE DROSOPHILA MELANOGASTER

Abstract. — Drosophila and other ectotherms show geographic genetic variation in body size, with larger individuals at higher latitudes and altitudes. Temperature is implicated as an important selective agent because long-term laboratory culture of Drosophila leads to the evolution of larger body size at lower temperatures. In this paper, we tested the hypothesis that, in Drosophila melanogaster, larger size is favored at lower temperatures in part because of selection on adult females. We used replicated lines of D. melanogaster artificially selected for increased and decreased wing area with constant cell area. The resulting size differences between the selected lines were due solely to differences in cell number, and thereby were similar to the cellular basis of clinal variation in body size in nature. We examined life-history traits of adult females at 18 and 25°C. Rearing for two generations at the two temperatures did not affect the extent of the size differences between lines from the different selection regimes. There was a strong interaction between temperature and size selection for both survival and lifetime reproductive success, with larger females living significantly longer and producing more offspring over their lifetime only when reared and tested in the colder environment. There was also an increase in average daily progeny production in large-line females relative to the control and small lines again, only in the colder environment. Thus, the females from the large selection lines were relatively fitter at the colder temperature. At both experimental temperatures, especially the lower one, the small- line females rescheduled their progeny production to later ages. Larger body size may have evolved at higher latitudes and altitudes because of the advantages to the adult female of being larger at lower temperatures.