Life history parameters and diet of Risso's dolphins,Grampus griseus,from southeastern South Africa

The life history of Risso's dolphins (Grampus griseus) remains poorly known and data from strandings can help provide important information. Data from 126 Risso's dolphins stranded or bycaught along the southeastern coastline of South Africa between 1958 and 2017 were analyzed in relation to their sex, age structure, and diet. Mean estimated length at birth was 146.9 cm, while maximum length was 325 cm for males and 313 cm for females; small sample sizes precluded detailed examination of sexual dimorphism. Age estimates for 33 individuals (14 males, 17 females, 2 unknown sex) indicated a maximum age of 13 years (males) and 17 years (females), respectively; the oldest animal was 19 years (unknown sex). Mean length and age at attainment of sexual maturity were estimated at 280 cm and 7.1 years in males and at 282 cm and 7.7 years in females. Stomach contents from 27 individuals showed that diets of immature and mature males and females overlapped and consisted predominantly of cephalopods. Reported strandings decreased between 2000 and 2017, possibly due to a lack of reporting associated with a ban on driving on beaches or related to the collapse of the local “chokka” squid (Loligo reynaudii) fishery in 2014–2015.     

Risso's dolphins alter daily resting pattern in response to whale watching at the Azores

Behavioral responses of Risso's dolphins (Grampus griseus) to whale watching vessels were studied off Pico Island, Azores. Dolphin behavior was studied from a land‐based lookout, enabling observations of groups in the absence and presence of vessels. The number of whale watching vessels showed a clear seasonal pattern, dividing the whale watching period into a low season and a high season. During the low season, Risso's dolphins rested mainly in the morning and afternoon. During the high season, Risso's dolphins rested less and did so mainly at noon, when the number of active vessels was lowest. Data analysis using a generalized additive mixed model indicated that this change in resting behavior was associated with vessel abundance. When more than five vessels were present, Risso's dolphins spent significantly less time resting and socializing. During the high season, this vessel abundance was exceeded during 20% of observation days. While we cannot be sure that the observed changes in behavior have fitness consequences for Risso's dolphins, reduced resting and socializing rates can have negative impacts on the build‐up of energy reserves and on reproductive success. We suggest the adoption of precautionary management measures to regulate the timing and intensity of whale watching activities.     

LIFE HISTORY OF FRASER'S DOLPHIN, LAGENODELPHIS HOSEI, BASED ON A SCHOOL CAPTURED OFF THE PACIFIC COAST OF JAPAN

Life history of Fraser's dolphin, a little known delphinid species, was examined based on 108 specimens from a school captured by the driving fishing method in Japan. The sex ratio was approximately 1:1, and mature dolphins of both sexes formed the bulk of the school. The oldest animals were two males and a female of 17.5 yr. Age and body length at sexual maturity were estimated at 7–10 yr and 220–230 cm in males and 5–8 yr and 210–220 cm in females. Mature males were larger in body length than mature females and showed apparent secondary sexual features: deepening of the tail stock and widening and darkening of the lateral dark stripe. The annual ovulation rate was 0.49. The estimated neonatal length (110 cm) predicts a gestation period of about 12.5 mo and calving peaks in spring and probably also in fall. The calving interval was estimated to be about 2 yr. These life history parameters are similar to those of the striped and pantropical spotted dolphins, except for longevity. The reproductive rate of this species may be lower than that of other pelagic delphinids, if the observed shorter longevity is real.     

Hip bone morphometrics of bottlenose dolphins (Tursiops truncatus) from Adriatic Sea: Sex determination and postnatal development

Bones are frequently the only cetacean samples available in the wild and may be useful for determination of sex, physical maturity, body length, and body mass. We analyzed the postnatal morphological development and morphometric characteristics of the cleaned and dried, paired hip bone (os coxae) from 131 bottlenose dolphins, Tursiops truncatus, (61 females, 70 males; total body length 117–322 cm) ranging in age from neonates to 28 yr, salvaged from Croatian waters of the eastern Adriatic Sea. Adult dolphins exhibited sexual dimorphism in their hip bone morphology and morphometrics: fully‐formed hip bones from males were significantly longer, heavier, and more robust than those from females. Moreover, the fully formed hip bone from males featured a prominent bony tuberosity protruding dorsolaterally from the caudal half that was significantly wider and higher than in females. Morphometric measurements were used to generate classification functions for reliable sex determination (97%). In addition, hip bone length in both sexes correlated tightly with total body length, allowing one measurement to be calculated from the other. Only a single ossification developed in all hip bones although soft tissue attachments and comparative review suggests homological equivalence with the tripartite terrestrial mammal hip bone.     

Global distribution of Risso's dolphin Grampus griseus: a review and critical evaluation

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Sexual maturity and estimated fecundity in female Indo‐Pacific bottlenose dolphins (Tursiops aduncus) from South Australia: Combining field observations and postmortem results

Knowledge of life history of bottlenose dolphins in Australia is limited, with no studies comparing field and postmortem data from the same region. Carcasses of 88 opportunistically collected females (1987–2015) were studied, including eight observed for up to 23 yr before death. Ovaries were weighed and number of corpora determined grossly. Nineteen mammary glands were examined histologically. Relative age was determined using developmental features and body length, estimated age by tooth incremental lines, and physical maturity by epiphyseal fusion. Age at sexual maturation was 7.17 yr (6–>14 yr) at body lengths of 191–209 cm. Physical maturation occurred at 12–17 yr, and body lengths of 195–233 cm. An Exponential model provided the best fit for age at body length, with an asymptote at 214 cm. Combined ovary weight increased between birth and sexual maturation, with significantly more corpora in the left ovary. Corpora increased with age and appeared to persist throughout life. For mature females, 95% had ≤ 11 corpora. Number of corpora was consistent with number of calves for females with known life histories. Estimated interbirth period was about 4 yr. Three females with abnormal reproductive features were either chronically diseased and/or had high heavy metal burdens.     

External morphology of the short-beaked common dolphin, Delphinus delphis: growth, allometric relationships and sexual dimorphism

Growth, allometric relationships and sexual dimorphism are described from measurements of 105 male, 149 female and 38 unsexed specimens of short‐beaked common dolphin, Delphinus delphis, stranded along the Irish coastline (53.8% of the sample) or by‐caught in fisheries (46.2% of the sample), from 1990 to 2003. For each dolphin, 24 external body length measurements were recorded. Ages were determined for 183 dolphins by analysis of growth layer groups in the dentine. Males ranged in total body length (TBL) from 105 to 231 cm and females from 93 to 230 cm, with a maximum age of 25 years obtained for both sexes. Using a single Gompertz growth curve, asymptotic values obtained for TBL were 211.6 cm and 197.4 cm for males and females, respectively. Asymptotic lengths were attained at 11 years in males and 9 years in females. The gestation period was estimated to last approximately 11.5 months. Sexual size dimorphism (SSD) was evident, with males being significantly larger than females for 20 of the characters measured, and an SSD ratio of 1.06 was obtained. Sexual shape dimorphism was lacking, except for the presence of prominent postanal humps in mature males.     

Annual variability in dentin δ15N and δ13C reveal sex differences in weaning age and feeding habits in Risso's dolphins (Grampus griseus)

Teeth of odontocetes accumulate annual dentinal growth layer groups (GLGs) that record isotope ratios, which reflect the time of their synthesis. Collectively, they provide lifetime records of individual feeding patterns from which life history traits can be inferred. We subsampled the prenatal dentin and postnatal GLGs in Risso's dolphins (Grampus griseus) (n = 65) that stranded or were collected as bycatch in Taiwan (1994–2014) and analyzed them for δ¹⁵N and δ¹³C. Age‐specific δ¹⁵N and δ¹³C values were corrected for effects of calendar year, stranding site, C/N, and sex. δ¹⁵N values were higher in prenatal layers (14.94‰ ± 0.74‰) than in adult female GLGs (12.58‰ ± 0.20‰), suggesting fetal enrichment during gestation. Decreasing δ¹⁵N values in early GLGs suggested changes in dietary protein sources during transition to complete weaning. Weaning age was earlier in males (1.09 yr) than in females (1.81 yr). Significant differences in δ¹⁵N values between weaned males and females suggest potential sexual segregation in feeding habits. δ¹³C values increased from the prenatal to the 4th GLG by ~1.0‰, indicative of a diet shift from ¹³C‐depleted milk to prey items. Our results provide novel insights into the sex‐specific ontogenetic changes in feeding patterns and some life history traits of Risso's dolphins.     

Life history of harbor porpoises (Phocoena phocoena) in Scottish (UK) waters

Life history parameters were determined for stranded and bycaught harbor porpoises (Phocoena phocoena) from Scottish (UK) waters (1992–2005). Fetal growth rate was 84.4 mm/mo and mean size at birth was 76.4 cm (range 65–88 cm). Males and females had a similar range of body lengths (65–170 cm and 66–173 cm, respectively), although asymptotic lengths were higher in females than males (approximately 158 cm and 147 cm, respectively). Nonpregnant females were significantly lighter, in relation to their length than males. Maximum estimated age was 20 yr for both sexes. Age at sexual maturity (ASM) was estimated as 4.35 yr in females and 5.00 yr in males. Conception occurred mainly in July and August although reproductively active males were recorded during April to July. Gestation lasted 10–11 mo, with calving mainly between May and July. Lactating females were recorded during June to November, while small calves with solid food in their stomachs were found mainly during February to May. Estimated pregnancy rate (0.34–0.40) is lower than recorded elsewhere, but is likely underestimated due to the prevalence of mature females of poor health status in the sample. Nevertheless, cetacean strandings can be an essential source of data on demographic parameters.     

Sex and reproduction of the alfonsino Beryx splendens (Pisces, Berycidae) from the Macaronesian archipelagos

This paper provides comparative information on the reproductive biology of the alfonsino, Beryx splendens Lowe, 1834, species with commercial interest in the Azores, Madeira and the Canary Islands. A total of 846 individuals from Azores (14.0–42.0 cm fork length), 621 from Madeira (17.2–50.0 cm fork length) and 643 from the Canaries (18.2–38.9 cm fork length) were used for the study. The alfonsino is gonochoric with no evidence of sexual dimorphism. Females are more abundant than males; this dominance probably reflects certain differences in the spatial distribution and/or the catchability of males and females in the Macaronesian archipelagos. The spawning season was distinct for the three Macaronesian areas, with an observed North–South variation in the reproductive period: September–March in the Azores, March–June in Madeira and July–September in the Canary Islands. The size at sexual maturity estimated for Madeira and the Canary Islands is similar (32 and 30 cm fork length, respectively), while for the Azores it is reached at smaller length (23 cm fork length). The differences observed in the size at sexual maturity can be explained by the different exploitation levels in each archipelago. Life‐history parameters of the alfonsino suggest that this species has a specialistic life‐history strategy and fisheries based on this species are more susceptible to growth overfishing and population depletion.     

Reproductive biology of female striped dolphin (Stenella coeruleoalba) from the western Mediterranean

Data on age and length at sexual maturity, calving interval, lactation period, and ovulation and pregnancy rates were analysed from 84 female striped dolphin collected during the Mediterranean morbillivirus epizootic. Age was determined by counting growth layer groups in tooth dentine, and reproductive status was determined by the macroscopic examination of the ovaries and the uterus. Age at sexual maturity (ASM) was estimated using two different methods; giving average estimates of 12 and 12.3 years. The average length at sexual maturity was 187 cm and the calving interval was four years. The annual pregnancy rate (APR) was calculated using two different criteria. In the first, only females that were pregnant at the time of death, as evidenced by the presence of a foetus, were incorporated in the calculations. This method gave an estimate of 8.9% as the apparent pregnancy rate of the dolphins killed by the epizootic. The second method incorporated in the calculations females that were pregnant at the time of death and females that had young corpus albicans in the ovaries and a distended uterus. This method is considered to reflect the true pregnancy rate of the population and gave an estimate of 25%. Estimates of ovulation rate (0.401) and lactation period (16 months) were considered to be potentially affected by the disease and, therefore, biased. These estimates reflect a peculiar situation of the diseased dolphins but are useful for comparative purposes. Estimates of ASM and APR, parameters which are not likely to be affected by the disease, differed markedly from those of other Stenella populations and could indicate a density-dependent response, possibly reflecting food limitations.     

AGE, GROWTH, AND REPRODUCTION OF THE FINLESS PORPOISE, NEOPHOCAENA PHOCAENOIDES, IN THE COASTAL WATERS OF WESTERN KYUSHU, JAPAN

Abstract: In the coastal waters of western Kyushu, Japan, a total of 97 incidentally taken or stranded finless porpoises, Neophocaena phocaenoides , was collected for studying age, growth and reproduction. An additional 17 specimens from the Inland Sea were used for a comparison of life history. Mean neonatal body length was 78.2 cm. Both males and females grew to around 140 cm by 5 yr of age. The maximum body lengths of males and females in western Kyushu were 174.5 cm and 165.0 cm, respectively, which were smaller than those recorded in other Japanese waters. Females probably attain sexual maturity at ages of 6–9 yr and at body lengths of 135–145 cm. Males probably mature sexually at ages of 4–6 yr, at body lengths of 135–140 cm and at weight of testis of 40–150 g. The lack of females aged 5–6 yr and males aged 4–5 yr precluded firm conclusions on ages at sexual maturity. Parturition in western Kyushu was estimated to be prolonged from autumn to spring, whereas in the Inland Sea and Pacific waters it was restricted from spring to summer with a peak in April. These geographical differences and available information on distribution implies that the finless porpoises in western Kyushu constitute a local population.     

Age and Growth of Blue Antimora <Emphasis Type="Italic">Antimora rostrata</Emphasis> (Moridae) in Southwestern Greenland Waters

The results of determination of age and study on growth of blue antimora Antimora rostrata from the waters of Southwestern Greenland are presented. The results are based on the analysis of 200 fish otoliths. In the catches, we found specimens of antimora with total lengths of 18?70 cm, body weights of 23?2731 g, at the age of 7?38 years. Minimal age of males (not considering juvenile individuals) was 10 years at body length of 27?33 cm; maximal age was 18 years at 42 cm. Minimal age of females was 9 years at length of 21–27 cm; maximal age was 38 years at 70 cm. The rate of linear growth in blue antimora from Southwestern Greenland waters is comparable to that in the fish from New Zealand and Ross Sea waters but considerably lower than indicated earlier for fish from the waters of Iceland, Greenland, and the Mid-Atlantic Ridge. The age of reaching sexual maturity in males and females is preliminary determined as 15 and 19?20 years, respectively.     

Reproductive biology of brown trout, Salmo trutta macrostigma Dumeril 1858, in a tributary of the Ceyhan River which flows into the eastern Mediterranean Sea

The study describes some key elements of the reproductive biology, including spawning season, age at sexual maturity, fecundity and egg diameter of the native brown trout, Salmo trutta macrostigma, in a tributary of the Ceyhan River. A total of 197 brown trout (118 females and 79 males) were captured in 2000–2001 by electric fishing. In observations on monthly changes, the gonadosomatic index (GSI) and the monthly frequency distribution of egg diameter confirmed that spawning lasted from November to January. Some 27.7% of the females and 62.5% of the males attained sexual maturity in their second year. The smallest fork length (FL) of brown trout attaining sexual maturity was 17.4 cm for males and 17.8 cm for females. Mean fecundity in age groups II, III, IV and V were 360, 452, 693 and 1283 eggs per female, respectively. One 9‐year‐old female had a unique 3232 egg count. The mean fecundity of the sampled population was 554 eggs per fish, positively correlated with the FL (mm) (R = 0.8227 ) and body weight (R = 0.8130). The diameter of mature eggs in the spawning season ranged from 3.250 to 5.930 mm, with a 4.146 mm average. Mean egg diameter in age groups II, III, IV and V in the spawning season were 0.813, 3.799, 4.663 and 5.243 mm, respectively. Fecundity, egg weight and diameter were statistically different in all age groups.     

Life history of the Indo‐Pacific humpback dolphin in the Pearl River Estuary,southern China

We studied life history characteristics of the Hong Kong/Pearl River Estuary population of Indo‐Pacific humpback dolphins (Sousa chinensis), based on data from 120 specimens stranded between 1995 and 2009, 40 individuals biopsied at sea, and a long‐term (14+ yr) photo‐identification study. Ages were determined for 112 specimens by thin‐sectioning teeth and counting growth layer groups. Estimated length at birth was 101 cm. Longevity was at least 38 yr, and there was little difference in growth patterns of males and females. Growth was described by a Bayesian two‐phase Gompertz model; asymptotic length was reached at 249 cm. The tooth pulp cavity filled at an average of 18.5 yr of age. Physical maturity was reached at between 14 and 17 yr of age, apparently a few years after attainment of sexual maturity. Maximum lengths and weights of about 268 cm and 240 kg were attained. Females appear to lose all their spots by 30 yr, although males may retain some spotting throughout life. Calving occurred throughout the year, with a broad peak from March to June. Of 60 females monitored at sea for >14 yr of the study, none were documented to have more than three calves, suggestive of low reproductive output or low calf survival.     

Patterns of growth in wild bottlenose dolphins, Tursiops truncatus

The growth of bottlenose dolphins is described from observations made during a capture release programme that has operated in coastal waters of the eastern Gulf of Mexico from 1970 to the present. Measurements of standard length, girth and body mass were recorded from 47 female and 49 male dolphins, some captured as many as nine times. Ages were known from approximate birth dates or estimated from counts of dentinal growth layers. In all three measurements. females grew at a faster initial rate than males, but reached asymptotic size at an earlier age. This extended period of growth in males resulted in significant sexual dimorphism in length, girth and mass at physical maturity. The growth of both sexes was well described by three-parameter Gompertz models using either cross-sectional data or a mixture of longitudinal and cross-sectional data. There was considerable variation in size-at-age for both sexes in all year classes. Residuals of size measurements were used to derive measures of relative size for individual dolphins; most dolphins demonstrated little ontogenetic change in relative size. Body mass was adequately predicted by multiple regression equations that incorporated both length and girth as independent variables.     

Latitudinal variation in sexual dimorphism in life‐history traits of a freshwater fish

Sexual dimorphism is common across the animal kingdom, but the contribution of environmental factors shaping differences between the sexes remains controversial. In ectotherms, life‐history traits are known to correlate with latitude, but sex‐specific responses are not well understood. We analyzed life‐history trait variation between the sexes of European perch (Perca fluviatilis L.), a common freshwater fish displaying larger female size, by employing a wide latitudinal gradient. We expected to find sex‐dependent latitudinal variation in life‐history variables: length at age, length increment, and size at maturity, with females showing consistently higher values than males at all latitudes. We further anticipated that this gender difference would progressively decrease with the increasingly harsh environmental conditions toward higher latitude. We hypothesized that growth and length increment would decrease and size/age at maturity would increase at higher latitudes. Our results confirmed female‐biased sexual size dimorphism at all latitudes and the magnitude of sexual dimorphism diminished with increase in latitude. Growth of both sexes decreased with increase in latitude, and the female latitudinal clines were steeper than those of males. Hence, we challenge two predominant ecological rules (Rensch's and Bergmann's rules) that describe common large‐scale patterns of body size variation. Our data demonstrate that these two rules are not universally applicable in ectotherms or female‐biased species. Our study highlights the importance of sex‐specific differences in life‐history traits along a latitudinal gradient, with evident implications for a wide range of studies from individual to ecosystems level.     

Odontocete occurrence in relation to changes in oceanography at a remote equatorial Pacific seamount

Seamounts are considered hot spots of biodiversity and can aggregate pelagic predators and their prey. Passive acoustic monitoring was conducted over 3 mo in 2012 to document the occurrence of odontocetes near a seamount chain in the central equatorial Pacific in relation to oceanographic changes over time. Beaked whale echolocation signals were most frequently encountered. The main beaked whale signal was an unknown type, BW38, which resembled signals produced by Blainville's beaked whales. It had high occurrence during high sea surface temperature and low sea surface salinity. Cuvier's beaked whales were the second most detected. They had an opposite pattern and were encountered more often when sea surface temperature was low and net primary productivity was high. Risso's dolphins and short‐finned pilot whales had high acoustic densities, and echolocated predominantly at night. Risso's dolphins occurred more often during low sea surface height deviation. False killer whales were less frequently detected and mostly occurred during the day. Sperm whale detections were fewer than expected and associated with high chlorophyll a. Short duration Kogiidae encounters occurred on average every third day. These types of long‐term site studies are an informative tool to comparatively assess species composition, relative abundance, and relationship to oceanographic changes.     

Evolution of biometric and life‐history traits in lizards (Gallotia) from the Canary Islands

The aim was to study as to how biometric and life‐history traits of endemic lacertids in the Canary Islands (genus Gallotia) may have evolved, and possible factors affecting the diversification process of this taxon on successively appearing islands have been deduced. To that end, comparative analyses of sexual dimorphism and scaling of different body, head and life‐history traits to body size in 10 species/subspecies of Gallotia have been carried out. Both Felsenstein's independent contrasts and Huey and Bennett's ‘minimum evolution’ analyses show that male and female snout‐vent length (SVL) changed proportionally (sexual size dimorphism not changing with body size) throughout the evolution of these lizards and all within‐sex biometric traits have changed proportionally to SVL. Life‐history traits (size at sexual maturity, clutch size, hatchling SVL and mass, and life span) are highly correlated with adult female body size, the first two being the only traits with a positive allometry to female SVL. These results, together with the finding that the slope of hatchling SVL to female SVL regression was lower than that of SVL at maturity to female SVL, indicates that larger females reach maturity at a larger size, have larger clutches and, at the same time, have relatively smaller hatchlings than smaller females. There was no significant correlation between any pair of life‐history traits after statistically removing the effect of body size. As most traits changed proportionally to SVL, the major evolutionary change has been that of body size (a ca. threefold change between the largest and the smallest species), that is suggested to be the effect of variable ecological conditions faced by founder lizards in each island.     

High calf mortality in bottlenose dolphins in the Bay of Islands,New Zealand–a local unit in decline

Bottlenose dolphins (Tursiops truncatus) in the Bay of Islands, New Zealand, have been studied for almost two decades. Since 2003, fewer than 150 dolphins visited the bay during each season and the local unit has declined 7.5% annually from 1997 to 2006. The causes of decline are unclear but probably include mortality and emigration. Here, we used a long‐term database to estimate reproductive parameters of female bottlenose dolphins including recruitment rates. A total of 704 surveys were conducted in which 5,577 sightings of 408 individually identified dolphins were collected; of these 53 individuals were identified as reproductive females. The calving rate increased between periods (1997–1999 = 0.13, CL = 0.07–0.21; 2003–2005 = 0.25, CL = 0.16–0.35 calves/reproductive female/year). A 0.25 calving rate suggests that on average, a female gives birth only once every four years, which is consistent with the estimated calving interval (4.3 yr, SD = 1.45) but still is lower than values reported for other populations. Conversely, apparent mortality rates to age 1+ (range: 0.34–0.52) and 2+ (range: 0.15–0.59) were higher than values reported elsewhere. The high apparent calf mortality in conjunction with a decline in local abundance, highlight the vulnerability of bottlenose dolphins in the Bay of Islands. Long‐term studies are required to understand the causes of high calf mortality and the decline in local abundance. Meanwhile, management should focus on minimizing sources of anthropogenic disturbance and enforcing compliance with current legislation.