Mariano Sironi

Isotopic and genetic evidence for culturally inherited site fidelity to feeding grounds in southern right whales (Eubalaena australis)

Ocean warming will undoubtedly affect the migratory patterns of many marine species, but specific changes can be predicted only where behavioural mechanisms guiding migration are understood. Southern right whales show maternally inherited site fidelity to near‐shore winter nursery grounds, but exactly where they feed in summer (collectively and individually) remains mysterious. They consume huge quantities of copepods and krill, and their reproductive rates respond to fluctuations in krill abundance linked to El Niño Southern Oscillation (ENSO). Here we show that genetic and isotopic signatures, analysed together, indicate maternally directed site fidelity to diverse summer feeding grounds for female right whales calving at Península Valdés, Argentina. Isotopic values from 131 skin samples span a broad range (–23.1 to –17.2‰δ13C, 6.0 to 13.8‰δ15N) and are more similar than expected among individuals sharing the same mitochondrial haplotype. This pattern indicates that calves learn summer feeding locations from their mothers, and that the timescale of culturally inherited site fidelity to feeding grounds is at least several generations. Such conservatism would be expected to limit the exploration of new feeding opportunities, and may explain why this population shows increased rates of reproductive failure in years following elevated sea‐surface temperature anomalies off South Georgia, the richest known feeding ground for baleen whales in the South Atlantic.

Population histories of right whales (Cetacea: Eubalaena) inferred from mitochondrial sequence diversities and divergences of their whale lice (Amphipoda: Cyamus)

Right whales carry large populations of three ‘whale lice’ (Cyamus ovalis, Cyamus gracilis, Cyamus erraticus) that have no other hosts. We used sequence variation in the mitochondrial COI gene to ask (i) whether cyamid population structures might reveal associations among right whale individuals and subpopulations, (ii) whether the divergences of the three nominally conspecific cyamid species on North Atlantic, North Pacific, and southern right whales (Eubalaena glacialis, Eubalaena japonica, Eubalaena australis) might indicate their times of separation, and (iii) whether the shapes of cyamid gene trees might contain information about changes in the population sizes of right whales. We found high levels of nucleotide diversity but almost no population structure within oceans, indicating large effective population sizes and high rates of transfer between whales and subpopulations. North Atlantic and Southern Ocean populations of all three species are reciprocally monophyletic, and North Pacific C. erraticus is well separated from North Atlantic and southern C. erraticus. Mitochondrial clock calibrations suggest that these divergences occurred around 6 million years ago (Ma), and that the Eubalaena mitochondrial clock is very slow. North Pacific C. ovalis forms a clade inside the southern C. ovalis gene tree, implying that at least one right whale has crossed the equator in the Pacific Ocean within the last 1–2 million years (Myr). Low‐frequency polymorphisms are more common than expected under neutrality for populations of constant size, but there is no obvious signal of rapid, interspecifically congruent expansion of the kind that would be expected if North Atlantic or southern right whales had experienced a prolonged population bottleneck within the last 0.5 Myr.

Increased Wounding of Southern Right Whale (Eubalaena australis) Calves by Kelp Gulls (Larus dominicanus) at Península Valdés, Argentina.

At least 626 southern right whale (Eubalaena australis) calves died at the Península Valdés calving ground, Argentina, between 2003 and 2014. Intense gull harassment may have contributed to these deaths. In the 1970s, Kelp Gulls (Larus dominicanus) began feeding on skin and blubber pecked from the backs of living right whales at Valdés. The frequency of gull attacks has increased dramatically over the last three decades and mother-calf pairs are the primary targets. Pairs attacked by gulls spend less time nursing, resting and playing than pairs not under attack. In successive attacks, gulls open new lesions on the whales’ backs or enlarge preexisting ones. Increased wounding could potentially lead to dehydration, impaired thermoregulation, and energy loss to wound healing. The presence, number and total area of gull-inflicted lesions were assessed using aerial survey photographs of living mother-calf pairs in 1974–2011 (n = 2680) and stranding photographs of dead calves (n = 192) in 2003–2011. The percentage of living mothers and calves with gull lesions increased from an average of 2% in the 1970s to 99% in the 2000s. In the 1980s and 1990s, mothers and calves had roughly equal numbers of lesions (one to five), but by the 2000s, calves had more lesions (nine or more) covering a greater area of their backs compared to their mothers. Living mother-calf pairs and dead calves in Golfo Nuevo had more lesions than those in Golfo San José in the 2000s. The number and area of lesions increased with calf age during the calving season. Intensified Kelp Gull harassment at Península Valdés could be compromising calf health and thereby contributing to the high average rate of calf mortality observed in recent years, but it cannot explain the large year-to-year variance in calf deaths since 2000.

Interannual Variation in the Stable Isotope Differences Between Mothers and Their Calves in Southern Right Whales (Eubalaena australis)

Lactation is the most energetically expensive aspect of mammalian reproduction. As capital breeders, lactating southern right whales (Eubalaena australis) are completely dependent on their stored nutrients. The relative proportion of different endogenous nutrient pools used during lactation could be assessed using stable isotopes. We determined the stable carbon and nitrogen isotope difference between skin samples of 42 southern right whale mothers and their calves. The mean δ 15 N value of calves was 0.51‰ higher than that of their mothers, but their δ 13 C values were identical. However, when analyzed by year, the mother-calf pairs showed no isotope differences in 2004, but calves had higher δ 15 N (0.85‰) and δ 13 C (0.63‰) in 2003 and 2005. We hypothesize that the interannual variability was a consequence of different levels of nutritional stress. A decline in food abundance prior to the nursing seasons could result in mothers with relatively poorer physical condition that would not be able to meet the high energetic demands of their offspring. Thus, the calves would be forced to utilize proteins as well as lipids to meet this demand, increasing their nitrogen and carbon isotope ratios. This hypothesis is supported by an independent assessment of the proportion of stranded whales over the same time period.

Metal Levels in Southern Right Whales ( Eubalaena australis ) from Península Valdés, Argentina

Península Valdes, Argentina, is a nursing ground for a population of southern right whales (Eubalaena australis). In the last two decades this area has been subjected to an increase in population, tourism and industrial growth. This has raised the concern for exposure to chemical contaminants such as metals. In this study we measured nonessential metals (Ag, Al, As, Au, Ba, Be, Co, Cd, Cr, Hg, Li, Ni, Pb, Sb, Sn, Sr, U and Ti), essential metals (Cu, Fe, Mg, Mn, Mo and Zn) and an essential element (Se) in skin biopsies from female southern right whales (n=10). This is the first report on tissue concentrations of metals in adult southern right whales. Overall, tissue values were on the low end of the spectrum and similar to the values reported in studies of mysticetes from other regions. Measured values do not reflect substantial amounts of accumulation and thus serve as a baseline.

Post-mortem findings in southern right whales Eubalaena australis at Península Valdés, Argentina, 2003-2012.

Between 2003 and 2012, 605 southern right whales (SRW; Eubalaena australis) were found dead along the shores of Península Valdés (PV), Argentina. These deaths included alarmingly high annual losses between 2007 and 2012, a peak number of deaths (116) in 2012, and a significant number of deaths across years in calves-of-the-year (544 of 605 [89.9%]; average = 60.4 yr(-1)). Post-mortem examination and pathogen testing were performed on 212 whales; 208 (98.1%) were calves-of-the-year and 48.0% of these were newborns or neonates. A known or probable cause of death was established in only a small number (6.6%) of cases. These included ship strike in a juvenile and blunt trauma or lacerations (n = 5), pneumonia (n = 4), myocarditis (n = 2), meningitis (n = 1), or myocarditis and meningitis (n = 1) in calves. Ante-mortem gull parasitism was the most common gross finding. It was associated with systemic disease in a single 1-2 mo old calf. Immunohistochemical labeling for canine distemper virus, Toxoplasma gondii and Brucella spp., and PCR for cetacean morbillivirus (CeMV), influenza A, and apicomplexan protozoa were negative on formalin-fixed, paraffin-embedded lung and brain samples from a subset of whales; PCR for Brucella spp. was positive in a newborn/neonate with pneumonia. Skin samples from whales with gull parasitism were PCR negative for CeMV, poxvirus, and papillomavirus. This is the first long-term study to investigate and summarize notable post-mortem findings in the PV SRW population. Consistent, significant findings within or between years to explain the majority of deaths and those in high-mortality years remain to be identified.

Lifetime glucocorticoid profiles in baleen of right whale calves: potential relationships to chronic stress of repeated wounding by Kelp Gulls

Baleen from stranded carcasses of five right whale calves (four southern and one North Atlantic) contain cortisol and corticosterone patterns that may represent longitudinal profiles of stress exposure over time. Calves with high wounding from Kelp Gulls had markedly higher glucocorticoids in postnatal baleen than did less wounded and unwounded calves.