Ancestor–descendant relationships in evolution: origin of the extant pygmy right whale,Caperea marginata

Ancestor–descendant relationships (ADRs), involving descent with modification, are the fundamental concept in evolution, but are usually difficult to recognize. We examined the cladistic relationship between the only reported fossil pygmy right whale, †Miocaperea pulchra, and its sole living relative, the enigmatic pygmy right whale Caperea marginata, the latter represented by both adult and juvenile specimens. †Miocaperea is phylogenetically bracketed between juvenile and adult Caperea marginata in morphologically based analyses, thus suggesting a possible ADR—the first so far identified within baleen whales (Cetacea: Mysticeti). The †Miocaperea–Caperea lineage may show long-term morphological stasis and, in turn, punctuated equilibrium.     

Phenological changes in North Atlantic right whale habitat use in Massachusetts Bay

The North Atlantic right whale (Eubalaena glacialis) is one of the world's most highly endangered baleen whales, with approximately 400–450 individuals remaining. Massachusetts Bay (MB) and Cape Cod Bay (CCB) together comprise one of seven areas in the Gulf of Maine where right whales seasonally congregate. Here, we report on acoustically detected presence of right whales in MB over a nearly 6 year period, July 2007–April 2013, a time of both rapid ocean warming throughout the Gulf of Maine and apparent changes in right whale migratory dynamics. We applied an automated detection algorithm to assess hourly presence of right whale “up‐calls” in recordings from a 19‐channel acoustic array covering approximately 4,000 km² in MB. Over the survey, up‐calls were detected in 95% of 8 day periods. In each year, as expected, we observed a “peak season” of elevated up‐call detections in late winter and early spring corresponding to the season when right whales congregate to feed in CCB. However, we also saw an increase in right whale occurrence during time periods thought to be part of the “off‐season.” With the exception of 2009–2010, when acoustic presence was unusually low, the mean percent of hours in which up‐calls were detected increased every year, both during the peak season (from 38% in 2008 to 70% in 2012), and during the summer–fall season (from 2% in 2007 to 13% in 2012). Over the entire study, the peak season start date varied between 17 January and 26 February. Changes in right whale phenology in MB likely reflect broadscale changes in habitat use in other areas within the species range. This study demonstrates the value of continuous long‐term survey datasets to detect and quantify shifts in cetacean habitat use as environmental conditions change and the long‐term continued survival of right whales remains uncertain.     

Comparative osteology and phylogenetic relationships of Miocaperea pulchra,the first fossil pygmy right whale genus and species (Cetacea,Mysticeti, Neobalaenidae)

A fossil pygmy right whale (Cetacea, Mysticeti, Neobalaenidae) with exquisitely preserved baleen is described for the first time in the history of cetacean palaeontology, providing a wealth of information about the evolutionary history and palaeobiogeography of Neobalaenidae. This exquisitely preserved specimen is assigned to a new genus and species, Miocaperea pulchra gen. et sp. nov. , and differs from Caperea marginata Gray, 1846, the only living taxon currently assigned to Neobalaenidae, in details of the temporal fossa and basicranium. A thorough comparative analysis of the skeleton of M. pulchra gen. et sp. nov. and C. marginata is also provided, and forms the basis of an extensive osteology‐based phylogenetic analysis, confirming the placement of M. pulchra gen. et sp. nov. within Neobalaenidae as well as the monophyly of Neobalaenidae and Balaenidae; the phylogenetic results support the validity of the superfamily Balaenoidea. No relationship with Balaenopteroidea was found by the present study, and thus the balaenopterid‐like morphological features observed in C. marginata must have resulted from parallel evolution. The presence of M. pulchra gen. et sp. nov. around 2000 km north from the northernmost sightings of C. marginata suggests that different ecological conditions were able to support pygmy right whale populations in what is now Peru, and that subsequent environmental change caused a southern shift in the distribution of the living neobalaenid whales. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 166 , 876–911.     

ABUNDANCE AND POPULATION TREND (1978-2001) OF WESTERN ARCTIC BOWHEAD WHALES SURVEYED NEAR BARROW, ALASKA

The 2001 survey of western Arctic (Bering, Chukchi, and Beaufort seas) bowhead whales was conducted from 5 April to 7 June near Barrow, Alaska. Visual observers recorded a total of 3,295 “new” (not seen before) and 532 “conditional” (possibly seen before) whales in 1,130 h of watch effort, including 121 new calves (3.7% of the new whales). Concurrent with the visual survey, passive acoustic surveillance was conducted almost continuously from 16 April to 31 May, resulting in 27,023 locations of vocalizing bowhead whales. The estimated number of whales within 4 km of the perch (N₄) was 7,025 (SE = 1,068). The estimated proportion of the whales within 4 km of the perch (P₄) was 0.862 (SE = 0.044, computed by a moving blocks bootstrap). Combining these, the abundance estimate (N₄/P₄) for 2001 is 10,470 (SE = 1, 351) with a 95% confidence interval of 8, 100–13, 500. The estimated annual rate of increase (ROI) of the population from 1978 to 2001 is 3.4% (95% CI 1.7%‐5%). Reports from hunters and results of an aerial survey in June 2001 indicate whales continued to pass Barrow after the survey had ended. In 2001 51% (572 h) of the watch was scored as occurring during “fair‐excellent” visibility conditions, somewhat lower than the average for all surveys since 1978. Sea ice in the leads and fog were the principle environmental factors affecting visibility for all years. The estimated rate of increase and the fact that the number of calves counted in 2001 is the highest ever recorded suggest a steady recovery of this population. Other populations of large balaenids, notably the North Atlantic right whale, have failed to recover despite 70 yr of protection. The recovery of the howhead whale is likely attributable to low anthropogenic mortality, a relatively pristine habitat, and a well‐managed subsistence hunt. Nonetheless, offshore oil development, increasing shipping traffic, changes in the Bering Sea ecosystem, sea ice retreat, and possibly killer whale predation within its range could impact this bowhead population and should be carefully monitored.     

Fight or flight: antipredator strategies of baleen whales

• 1 The significance of killer whale Orcinus orca predation on baleen whales (Mysticeti) has been a topic of considerable discussion and debate in recent years. Discourse has been constrained by poor understanding of predator‐prey dynamics, including the relative vulnerability of different mysticete species and age classes to killer whales and how these prey animals avoid predation. Here we provide an overview and analysis of predatory interactions between killer whales and mysticetes, with an emphasis on patterns of antipredator responses.
• 2 Responses of baleen whales to predatory advances and attacks by killer whales appear to fall into two distinct categories, which we term the fight and flight strategies. The fight strategy consists of active physical defence, including self‐defence by single individuals, defence of calves by their mothers and coordinated defence by groups of whales. It is documented for five mysticetes: southern right whale Eubalaena australis, North Atlantic right whale Eubalaena glacialis, bowhead whale Balaena mysticetus, humpback whale Megaptera novaeangliae and grey whale Eschrichtius robustus. The flight strategy consists of rapid (20–40 km/h) directional swimming away from killer whales and, if overtaken and attacked, individuals do little to defend themselves. This strategy is documented for six species in the genus Balaenoptera.
• 3 Many aspects of the life history, behaviour and morphology of mysticetes are consistent with their antipredator strategy, and we propose that evolution of these traits has been shaped by selection for reduced predation. Fight species tend to have robust body shapes and are slow but relatively manoeuvrable swimmers. They often calve or migrate in coastal areas where proximity to shallow water provides refuge and an advantage in defence. Most fight species have either callosities (rough and hardened patches of skin) or encrustations of barnacles on their bodies, which may serve (either primarily or secondarily) as weapons or armour for defence. Flight species have streamlined body shapes for high‐speed swimming and they can sustain speeds necessary to outrun pursuing killer whales (>15–20 km/h). These species tend to favour pelagic habitats and calving grounds where prolonged escape sprints from killer whales are possible.
• 4 The rarity of observed successful attacks by killer whales on baleen whales, especially adults, may be an indication of the effectiveness of these antipredator strategies. Baleen whales likely offer low profitability to killer whales, relative to some other marine mammal prey. High‐speed pursuit of flight species has a high energetic cost and a low probability of success while attacks on fight species can involve prolonged handling times and a risk of serious injury.

     

Southern right whale (Eubalaena australis) calf mortality at Península Valdés,Argentina: Are harmful algal blooms to blame?

Península Valdés (PV) in Argentina is an important calving ground for southern right whales (SRWs, Eubalaena australis). Since 2005, right whale mortality has increased at PV, with most of the deaths (~90%) being calves <3 mo old. We investigated the potential involvement of harmful algal blooms (HABs) in these deaths by examining data that include: timing of the SRW deaths, biotoxins in samples from dead SRWs, abundances of the diatom, Pseudo‐nitzschia spp., and the dinoflagellate, Alexandrium tamarense, shellfish harvesting closure dates, seasonal availability of whale prey at PV and satellite chlorophyll data. Evidence of the whales' exposure to HAB toxins includes trace levels of paralytic shellfish toxins (PSTs) and domoic acid (DA) in tissues of some dead whales, and fragments of Pseudo‐nitzschia spp. frustules in whale feces. Additionally, whales are present at PV during both closures of the shellfish industry (due to high levels of PSTs) and periods with high levels of Pseudo‐nitzschia spp. and A. tamarense. There is a positive statistical relationship between monthly Pseudo‐nitzschia densities (but not A. tamarense) and calf deaths in both gulfs of PV.     

Passive acoustic monitoring predicts daily variation in North Atlantic right whale presence and relative abundance in Roseway Basin,Canada

North Atlantic right whale monitoring in Roseway Basin, Canada, is primarily based on short‐term (<14 d) visual surveys conducted during August–September. Variability in survey effort has been the biggest limiting factor to studying changes in the population's occurrence and habitat use. Such efforts could be enhanced considerably using passive acoustic monitoring (PAM). We sought to determine if variation in whale presence, relative abundance, demography, and/or behavior (estimated through visual surveys) could be explained by variation in three right whale call types in this habitat. A generalized linear model was fit to 23 d of concurrent PAM and visual monitoring during four summers within the Roseway Basin Right Whale Critical Habitat boundaries. The model revealed significant positive relationships between relative abundance, call counts and presence of surface‐active group behavior. PAM can refine daily right whale presence estimates. While visual observations (n = 23 d) implied a 40% decline in right whale presence during 2014–2015 relative to 2004–2005, PAM data (n = 211 d) showed right whales were present between 71%–85% of survey days throughout all years analyzed. We demonstrate that PAM is a useful tool to extend periods of right whale monitoring, especially in areas where visual monitoring efforts may be limited.     

Visual and acoustic surveys for North Atlantic right whales,Eubalaena glacialis,in Cape Cod Bay,Massachusetts, 2001–2005: Management implications

North Atlantic right whales, Eubalaena glacialis, remain endangered, primarily due to excessive anthropogenic mortality. Current management protocols in US waters are triggered by identifying the presence of at least one right whale in a management area. We assessed whether acoustic detection of right whale contact calls can work as an alternative to visual aerial surveys for establishing their presence. Aerial survey and acoustic monitoring were conducted in Cape Cod Bay, Massachusetts, in 2001–2005 and used to evaluate and compare right whale detections. Over the 58 d with simultaneous aerial and acoustic coverage, aerial surveys saw whales on approximately two-thirds of the days during which acoustic monitoring heard whales. There was no strong relationship between numbers of whales seen during aerial surveys and numbers of contact calls detected on survey days. Results indicate acoustic monitoring is a more reliable mechanism than aerial survey for detecting right whales. Because simple detection is sufficient to trigger current management protocols, continuous, autonomous acoustic monitoring provides information of immediate management utility more reliably than aerial surveillance. Aerial surveys are still required to provide data for estimating population parameters and for visually assessing the frequency and severity of injuries from shipping and fishing and detecting injured and entangled right whales.     

NORTH ATLANTIC RIGHT WHALE DISTRIBUTION IN RELATION TO SEA-SURFACE TEMPERATURE IN THE SOUTHEASTERN UNITED STATES CALVING GROUNDS

Standardized aerial surveys were used to document the winter (December–March) distribution of North Atlantic right whales in their calving area off the coasts of Georgia and northeastern Florida (1991–1998). Survey data were collected within four survey zones in and adjacent to federally designated critical habitat. These data, including whale‐sighting locations and sampling effort, were used to describe right whale distribution in relation to sea‐surface temperature (SST) from satellite‐derived images. Locations where whales were sighted (n= 609) had an overall mean SST of 14.3°C ± 2.1° (range 8°–22°C). Data from two survey zones having sufficient data (including the “early warning system” (EWS) zone and the Florida nearshore) were pooled by season and stratified by month to investigate changes in monthly ambient SST and fine‐scale distribution patterns of right whales in relation to SST within spatially explicit search areas. Using Monte Carlo techniques, SSTs and latitudes (means and standard deviations) of locations where whales were sighted were compared to a sampling distribution of each variable derived from daily‐search areas. Overall, results support a nonrandom distribution of right whales in relation to SST: during resident months (January and February), whales exhibited low variability in observed SST and a suggested southward shift in whale distribution toward warmer SSTs in the EWS zone; while in the relatively warmer and southernmost survey zone (Florida nearshore), right whales were concentrated in the northern, cooler portion. Our results support that warm Gulf Stream waters, generally found south and east of delineated critical habitat, represent a thermal limit for right whales and play an important role in their distribution within the calving grounds. These results affirm the inclusion of SST in a multivariate predictive model for right whale distribution in their southeastern habitat.     

EXPERIMENTAL MODELING OF LARGE WHALE ENTANGLEMENT INJURIES

The abrasive impact of commercial fishing gear on right whale fluke tissue was modeled using a reciprocating load generator. A line passing over the leading edge of a fluke specimen was fixed at one end while the other end was loaded in an oscillatory manner. A 24‐h abrasion test using an accelerated loading rate of 60 cycles/min and a 9.0‐kg load on the fixed end of the most abrasive line tested (old sink line) was used to simulate a whale swimming 2 m/s towing four standard lobster buoys for five days. A tension load of 267 N was generated on the oscillatory side of the fluke. The test line failed to break the skin on the fluke but, instead, produced a compression furrow (maximum depth 0.31 cm) that closely resembled marks found on stranded right (Eubalaena glacialis) and humpback (Megaptera novaeangliae) whales. Different line types (float vs. sink) and ages of rope (new vs. old) produced furrows of varying depths and appearances. Further tests characterizing whale skin/fishing line interactions will provide a better understanding of the mechanisms that govern entanglement injuries and provide a framework for better forensic analysis and interpretation of gear‐related injuries observed in large whales.     

The systematics of right whales (Mysticeti: Balaenidae)

Balaenidae (right whales) are large, critically endangered baleen whales represented by four living species. The evolutionary relationships of balaenids are poorly known, with the number of genera, relationships to fossil taxa, and position within Mysticeti in contention. This study employs a comprehensive set of morphological characters to address aspects of balaenid phylogeny. A sister‐group relationship between neobalaenids and balaenids is strongly supported, although this conflicts with molecular evidence, which may be an artifact of long‐branch attraction (LBA). Monophyly of Balaenidae is supported, and three major clades are recognized: (1) extinct genus Balaenula, (2) extant and extinct species of the genus Eubalaena, and (3) extant and extinct species of the genus Balaena plus the extinct taxon, Balaenella. The relationships of these clades to one another, as well as to the early Miocene stem balaenid, Morenocetus parvus, remain unresolved. Pliocene taxa, Balaenula astensis and Balaenula balaenopsis, form a clade that is the sister group to the Japanese Pliocene Balaenula sp. Eubalaena glacialis and Pliocene Eubalaena belgica, are in an unresolved polytomy with a clade including E. japonica and E. australis. Extant and fossil species of Balaena form a monophyletic group that is sister group to the Dutch Pliocene Balaenella, although phylogenetic relationships within Balaena remain unresolved.     

The oldest marine vertebrate fossil from the volcanic island of Iceland: a partial right whale skull from the high latitude Pliocene Tjörnes Formation

Extant baleen whales (Cetacea, Mysticeti) are a disparate and species‐rich group, but little is known about their fossil record in the northernmost Atlantic Ocean, a region that supports considerable extant cetacean diversity. Iceland's geographical setting, dividing North Atlantic and Arctic waters, renders it ideally situated to shed light on cetacean evolution in this region. However, as a volcanic island, Iceland exhibits very little marine sedimentary exposure, and fossil whales from Iceland older than the late Pleistocene are virtually unknown. Here, we present the first fossil whale found in situ from the Pliocene Tjörnes Formation (c. 4.5 Ma), Iceland's only substantial marine sedimentary outcrop. The specimen is diagnosed as a partial skull from a large right whale (Mysticeti, Balaenidae). This discovery highlights the Tjörnes Formation as a potentially productive fossil vertebrate locality. Additionally, this find indicates that right whales (Eubalaena) and bowhead whales (Balaena) were sympatric, with broadly overlapping latitudinal ranges in the Pliocene, in contrast to the modern latitudinal separation of their living counterparts.     

Global distribution of fin whales Balaenoptera physalus in the post‐whaling era (1980–2012)

相似文献   

Analysis of the C4 genes in baleen whales using a human cDNA probe

We have used a human C4 cDNA probe to investigate the complement component C4 gene in four members of the family Balaenopteridae: fin whale (Balaenoptera physalus), sei whale (B. borealis), minke whale (B. acutorostrata), and bryde's whale (B. edeni). Restriction mapping of genomic DNA from the first three species suggests the presence of only one locus in these species, and also shows that the C4 genes in the three species are very similar. We have used 14 restriction endonucleases to investigate the restriction fragment length polymorphism (RFLP) of fin whales, 13 enzymes for sei whales, and 8 enzymes for the minke whale. No polymorphism was seen in DNA from the five minke whale samples, but Rsa I and Taq I restriction enzymes gave polymorphism in fin and sei whales whereas Hind III and Msp I restriction enzymes showed polymorphism in sei whales only. Only one bryde's whale sample was available for investigation. The study of DNA available from mother-fetus pairs from the two polymorphic species demonstrated a simple, two-allele transmission of RFLP alleles.     

Association between pygmy right whales (Caperea marginata) and areas of high marine productivity off Australia and New Zealand

Abstract

Opportunistic sightings and strandings of Caperea marginata (n=196) from the vicinity of Australia and New Zealand (1884 to early 2007) were used to relate geographic and temporal patterns to oceanographic and broad-scale climatic variability. Records were not uniformly distributed along the coast and more (69%) were from Australia than New Zealand. Seven coastal whale ‘hotspots’ were identified which accounted for 61% of records with locality data. Half of the hotspot records were from southeast (37) and northwest (20) Tasmania—others each had 9–15 events. Upwelling and/or high zooplankton abundance has been documented near all whale hotspots. Records of C. marginata occurred in all months, with 75% in spring and summer. Inter-annual variability showed broad agreement between increased whale records (usually in spring/summer) and strongly positive ‘Niño 3.4’ during 1980–1995 but not thereafter. Coastal upwelling and productivity increase during climatic phenomena such as El Niño and are likely to be quickly beneficial to plankton-feeding whales such as C. marginata.     

Behaviour and kinematics of continuous ram filtration in bowhead whales (Balaena mysticetus)

Balaenid whales perform long breath-hold foraging dives despite a high drag from their ram filtration of zooplankton. To maximize the volume of prey acquired in a dive with limited oxygen supplies, balaenids must either filter feed only occasionally when prey density is particularly high, or they must swim at slow speeds while filtering to reduce drag and oxygen consumption. Using digital tags with three-axis accelerometers, we studied bowhead whales feeding off West Greenland and present here, to our knowledge, the first detailed data on the kinematics and swimming behaviour of a balaenid whale filter feeding at depth. Bowhead whales employ a continuous fluking gait throughout the bottom phase of foraging dives, moving at very slow speeds (less than 1 m s⁻¹), allowing them to filter feed continuously at depth. Despite the slow speeds, the large mouth aperture provides a water filtration rate of approximately 3 m³ s⁻¹, amounting to some 2000 tonnes of water and prey filtered per dive. We conclude that a food niche of dense, slow-moving zooplankton prey has led balaenids to evolve locomotor and filtering systems adapted to work against a high drag at swimming speeds of less than 0.07 body length s⁻¹ using a continuous fluking gait very different from that of nekton-feeding, aquatic predators.     

Rorqual whale (Balaenopteridae) surface lunge‐feeding behaviors: Standardized classification,repertoire diversity,and evolutionary analyses

Rorqual whales (Family: Balaenopteridae) are the world's largest predators and sometimes feed near or at the sea surface on small schooling prey. Most rorquals capture prey using a behavioral process known as lunge‐feeding that, when occurring at the surface, often exposes the mouth and head above the water. New technology has recently improved historical misconceptions about the natural variation in rorqual lunge‐feeding behavior yet missing from the literature is a dedicated study of the identification, use, and evolution of these behaviors when used to capture prey at the surface. Here we present results from a long‐term investigation of three rorqual whale species (minke whale, Balaenoptera acutorostrata; fin whale, B. physalus; and blue whale, B. musculus) that helped us develop a standardized classification system of surface lunge‐feeding (SLF) behaviors. We then tested for differences in frequency of these behaviors among the three species and across all rorqual species. Our results: (1) propose a unified classification system of six homologous SLF behaviors used by all living rorqual whale species; (2) demonstrate statistically significant differences in the frequency of each behavior by minke, fin, and blue whales; and (3) provide new information regarding the evolution of lunge‐feeding behaviors among rorqual whales.     

The fin whale Balaenoptera physalus (L. 1758) in the Mediterranean Sea

1. The ecology and status of fin whales Balaenoptera physalus in the Mediterranean Sea is reviewed. The species’ presence, morphology, distribution, movements, population structure, ecology and behaviour in this semi‐enclosed marine region are summarized, and the review is complemented with original, previously unpublished data. 2. Although the total size of the fin whale population in the Mediterranean is unknown, an estimate for a portion of the western basin, where most of the whales are known to live, was approximately 3500 individuals. High whale densities, comparable to those found in rich oceanic habitats, were found in well‐defined areas of high productivity. Most whales concentrate in the Ligurian‐Corsican‐Provençal Basin, where their presence is particularly noticeable during summer; however, neither their movement patterns throughout the region nor their seasonal cycle are clear. 3. Based on genetic studies, fin whales from the Mediterranean Sea are distinct from North Atlantic conspecifics, and may constitute a resident population, separate from those of the North Atlantic, despite the species’ historical presence in the Strait of Gibraltar. Fin whales are known to calve in the Mediterranean, with births peaking in November but occurring at lower rates throughout the year. They feed primarily on krill Meganyctiphanes norvegica which they capture by diving to depths in excess of 470 m. It is suggested that the extensive vertical migratory behaviour of its main prey may have influenced the social ecology of this population. 4. Known causes of mortality and threats, including collisions with vessels, entanglement in fishing gear, deliberate killing, disturbance, pollution and disease, are listed and discussed in view of the implementation of appropriate conservation measures to ensure the species’ survival in the region.     

SOUNDS OF A PYGMY RIGHT WHALE (CAPEREA MARGINATA)

Abstract: This is the first report of recordings of sounds from the pygmy right whale, Caperea marginata . The recordings were obtained in the presence of a juvenile in the harbor at Portland, on the southeastern corner of the Australian continent. Only one type of sound was heard-a short thump-like pulse or tone burst with a downsweep in frequency and decaying amplitude, with most energy between 60 and 120 Hz. The pulses occurred predominantly in pairs and once in a trio. The sounds are simpler than those of most baleen whales, but they show some similarity in characteristics. Source levels are in the lower end of the range determined for other species. There was no evidence to indicate the purpose of these sounds.