Molecular genetic identification of southern hemisphere beaked whales (Cetacea: Ziphiidae)

To assist in the species-level identification of stranded and hunted beaked whales, we compiled a database of 'reference' sequences from the mitochondrial DNA control region for 15 of the 20 described ziphiid species. Reference samples for eight species were obtained from stranded animals in New Zealand and South Australia. Sequences for a further seven species were obtained from a previously published report. This database was used to identify 20 'test' samples obtained from incompletely documented strandings around New Zealand. Analyses showed that four of these 'test' specimens (20%) had initially been misidentified. These included two animals of particular interest: (i) a Blainville's beaked whale ( Mesoplodon densirostris) , the first record of this species in New Zealand waters; and, (ii) a juvenile Andrews' beaked whale ( Mesoplodon bowdoini ), a species known from just over 20 strandings worldwide. A published sequence from a beaked whale product purchased in the Republic of Korea was identified as a Cuvier's beaked whale ( Ziphius cavirostris ). Levels of intra- and interspecific variation were compared to determine the potential for misidentification when the database or taxonomy is incomplete. Intraspecific variation was generally <2%, and interspecific divergence was generally >4.7%. Exceptions were within-species variation in Hyperoodon planifrons , southern bottlenosed whale (4.12%), which exceeded the variation between the two species of Berardius (3.78%), and variation between the two specimens assigned to M. hectori , Hector's beaked whale (7.14%). The latter case appears to be an error in species identification, and could represent the discovery of a new species of beaked whale.     

Geographic distribution of the Cross Seamount beaked whale based on acoustic detections

Beaked whales produce frequency-modulated echolocation pulses that appear to be species-specific, allowing passive acoustic monitoring to play a role in understanding spatio-temporal patterns. The Cross Seamount beaked whale is known only from its unique echolocation signal (BWC) with no confirmed species identification. This beaked whale spans the Pacific Ocean from the Mariana Archipelago to Baja California, Mexico, south to the equator, but only as far north as latitude 29°N. Within these warm waters, 92% of BWC detections occurred at night, 6% during crepuscular periods, and only 2% during daylight hours. Detections of BWC signals on drifting recorders with a vertical hydrophone array at 150 m depth demonstrated that foraging often occurred shallow in the water column (<150 m). No other species of beaked whale to date has been documented foraging in waters this shallow. Given their nocturnal, shallow foraging dives, this species appears to prefer prey that may be available in the water column only during those hours. The foraging behavior of Cross Seamount beaked whales appears to be unique among all beaked whales, and these findings contribute additional ecological and acoustic information which can help guide future efforts to identify this cryptic whale.     

中国大陆海域沿岸搁浅和误捕的喙鲸类调查

从2009 年1 月至2010 年2 月,我们对中国大陆海域沿岸搁浅和误捕的喙鲸类头骨标本进行了测量和鉴
定。目前中国大陆海域搁浅和误捕的喙鲸类有5 种,分别为柯氏喙鲸、贝氏喙鲸、朗氏喙鲸、柏氏中喙鲸和小
中喙鲸,其中小中喙鲸为中国海域(包括台湾)的首次报道,也是在西北太平洋区的首次记录。小中喙鲸的主
要形态鉴别特征为有1 对纺锤形牙齿着生在下颌骨联合后面。经对已报道过的大陆海域搁浅和误捕的3 个雌性
银杏齿中喙鲸头骨标本和台湾省搁浅银杏齿中喙鲸的颅骨、鼻骨、下颌骨以及牙齿特征对比重新鉴定,订正大
陆的3 个样本为雌性柏氏中喙鲸。本文报道了大陆沿岸海域搁浅和误捕的喙鲸类最新调查研究结果。     

APPEARANCE, DISTRIBUTION, AND GENETIC DISTINCTIVENESS OF LONGMAN'S BEAKED WHALE, INDOPACETUS PACIFICUS

Longman's beaked whale, Indopacetus pacificus , was known previously from only two skulls. Here we describe four new specimens of this species from strandings in the western and central Indian Ocean. Two juveniles, previously misidentified from external morphology as Hyperoodon planifrons , were identified as I. pacificus through diagnostic characteristics of mitochondrial (mt) DNA sequences derived from the holotype of this species. Images of the external appearance and teeth of the species are presented for the first time. Comparison of the color pattern of these new specimens with that of "tropical bottlenose whales" sighted in the tropical Indian and Pacific oceans confirm that those unidentified whales represent I. pacificus. Moore (1968) erected a new genus, Indopacetus , for this species (described initially as Mesoplodon pacificus ) based primarily on cranial morphology. Phylogenetic analyses of short mtDNA fragments available from the specimens known to date were unable to resolve the validity of this genus. However, the diagnostic osteological features highlighted by Moore (1968) for Indopacetus were also observed in the new specimens. Rib count and number of fused cervical vertebrae may also be diagnostic. Rostrum depth at mid-length and melon shape further distinguish this species from Mesoplodon beaked whales. As such, we see no reason on morphological grounds to overturn Moore's (1968) proposal that Longman's beaked whale is sufficiently distinct to be afforded its own genus.     

Stejneger's beaked whale strandings in Alaska, 1995–2020

Presented here is the first comprehensive and updated compilation of history, distribution, and findings of Stejneger's beaked whales (Mesoplodon stejnegeri) in Alaska. Stejneger's beaked whales are a poorly understood, elusive, deep-diving cetacean species found in the North Pacific Ocean. Since Stejneger's beaked whale strandings data in Alaska through 1994 were last published, 35 additional strandings have been documented. Twenty-seven animals stranded in the Aleutian Islands, seven stranded in Southcentral Alaska, and one animal stranded on St. Lawrence Island. Twenty-two carcasses were necropsied, but only four were fresh. Seventeen of the 22 died during mass stranding events and cause of death could not be definitively determined. Barotrauma was suspected in three cases and infectious disease possibly complicated by barotrauma occurred in two cases. We documented an expansion of strandings into the northern Bering Sea, characterized a sex bias, examined stomach contents that included macroplastic, and identified parasites not previously associated with Stejneger's beaked whales. Also included are data on the largest known mass stranding of Stejneger's beaked whales, which occurred on Adak Island in 2018. The history, distribution, and findings presented here are central to further our understanding of this species.     

CATCHES OF HUMPBACK AND OTHER WHALES FROM SHORE STATIONS AT MOSS LANDING AND TRINIDAD, CALIFORNIA, 1919–1926

Logbook data from California shore whaling stations at Moss Landing (1919–1922 and 1924) and Trinidad (1920 and 1922–1926) are analyzed. The logs for the two stations record the taking of 2,111 whales, including 1,871 humpbacks, 177 fin whales, 26 sei whales, 3 blue whales, 12 sperm whales, 7 gray whales, 1 right whale, 1 Baird's beaked whale, and 13 whales of unspecified type (probably humpbacks). Most whales were taken from spring to autumn, but catches were made in all months of some years. The sex ratios of humpback, fin, and sei whales (the three species with sufficient sample sizes to test) did not differ from parity. Primary prey, determined from stomach contents, included sardines and euphausiids for both humpback and fin whales, and 'plankton' (probably euphausiids) for sei whales. The prevalence of pregnancy was 0.46 among mature female humpbacks and 0.43 among mature female fin whales, although these values are reported with caution. Information on length distribution for all species is summarized. Analysis of the catch data for this and other areas supports the current view that humpback whales along the west coast of the continental United States comprise a single feeding stock and also suggests that the present population is well below pre-exploitation levels.     

Genetic structure of the beaked whale genus Berardius in the North Pacific,with genetic evidence for a new species

There are two recognized species in the genus Berardius, Baird's and Arnoux's beaked whales. In Japan, whalers have traditionally recognized two forms of Baird's beaked whales, the common “slate‐gray” form and a smaller, rare “black” form. Previous comparison of mtDNA control region sequences from three black specimens to gray specimens around Japan indicated that the two forms comprise different stocks and potentially different species. We have expanded sampling to include control region haplotypes of 178 Baird's beaked whales from across their range in the North Pacific. We identified five additional specimens of the black form from the Aleutian Islands and Bering Sea, for a total of eight “black” specimens. The divergence between mtDNA haplotypes of the black and gray forms of Baird's beaked whale was greater than their divergence from the congeneric Arnoux's beaked whale found in the Southern Ocean, and similar to that observed among other congeneric beaked whale species. Taken together, genetic evidence from specimens in Japan and across the North Pacific, combined with evidence of smaller adult body size, indicate presence of an unnamed species of Berardius in the North Pacific.     

Declining Abundance of Beaked Whales (Family Ziphiidae) in the California Current Large Marine Ecosystem

Beaked whales are among the most diverse yet least understood groups of marine mammals. A diverse set of mostly anthropogenic threats necessitates improvement in our ability to assess population status for this cryptic group. The Southwest Fisheries Science Center (NOAA) conducted six ship line-transect cetacean abundance surveys in the California Current off the contiguous western United States between 1991 and 2008. We used a Bayesian hidden-process modeling approach to estimate abundance and population trends of beaked whales using sightings data from these surveys. We also compiled records of beaked whale stranding events (3 genera, at least 8 species) on adjacent beaches from 1900 to 2012, to help assess population status of beaked whales in the northern part of the California Current. Bayesian posterior summaries for trend parameters provide strong evidence of declining beaked whale abundance in the study area. The probability of negative trend for Cuvier''s beaked whale (Ziphius cavirostris) during 1991–2008 was 0.84, with 1991 and 2008 estimates of 10771 (CV = 0.51) and ≈7550 (CV = 0.55), respectively. The probability of decline for Mesoplodon spp. (pooled across species) was 0.96, with 1991 and 2008 estimates of 2206 (CV = 0.46) and 811 (CV = 0.65). The mean posterior estimates for average rate of decline were 2.9% and 7.0% per year. There was no evidence of abundance trend for Baird''s beaked whale (Berardius bairdii), for which annual abundance estimates in the survey area ranged from ≈900 to 1300 (CV≈1.3). Stranding data were consistent with the survey results. Causes of apparent declines are unknown. Direct impacts of fisheries (bycatch) can be ruled out, but impacts of anthropogenic sound (e.g., naval active sonar) and ecosystem change are plausible hypotheses that merit investigation.     

SITE FIDELITY, ASSOCIATIONS, AND MOVEMENTS OF CUVIER'S (ZIPHIUS CAVIROSTRIS) AND BLAINVILLE'S (MESOPLODON DENSIROSTRIS) BEAKED WHALES OFF THE ISLAND OF HAWAI'I

Although the Ziphiidae are the second-most speciose family of cetaceans, information on beaked whale species and populations has been limited by the difficulties in finding and approaching free-ranging individuals. Site fidelity, patterns of association, and movements of two species, Cuvier's ( Ziphius cavirostris ) and Blainville's ( Mesoplodon densirostris ) beaked whales, were assessed using a 21-yr photographic data set from the west coast of the island of Hawai i. Resightings of individuals of both species spanned 15 yr, suggesting long-term site fidelity to the area. Long-term resightings were documented primarily from adult females of both species. Group sizes for both species were small and most groups had only a single adult male present. For Blainville's beaked whales, repeated associations between adult females and adult males were documented for all resightings of adult males over periods from 1 to 154 d. Among adult females, although repeated associations occurred up to 9 yr apart, individuals were seen separately in intervening years. Individuals of both species seen on multiple occasions were typically documented in multiple months/seasons, suggesting they may use the study area throughout the year. Such long-term site fidelity has implications both for potential population structure and for susceptibility of beaked whale populations to anthropogenic impacts.     

CHARACTERIZATION OF BEAKED WHALE (ZIPHIIDAE) AND SPERM WHALE (PHYSETER MACROCEPHALUS) SUMMER HABITAT IN SHELF-EDGE AND DEEPER WATERS OFF THE NORTHEAST U. S.

Sperm whales ( Physeter macrocephalus ) and beaked whales ( Mesoplodon spp. and Ziphius cavirostris ) are deep-diving cetaceans that frequent shelf-edge and Gulf Stream waters off the northeast U. S. coast. Sighting data collected during seven summer (1990, 1991, 1993, and 1995–1998) shipboard surveys were analyzed using a geographic information system to determine habitat use based on bathymetric and oceanographic features. Although sighting rates were lower for beaked whales, both taxa occupied similar habitats. Beaked whales were concentrated at the colder shelf edge, whereas sperm whales were associated with warmer off-shelf water. Mean sighting rates for both taxa were higher in canyon features, but only beaked whale sighting rates were significantly different between canyon and non-canyon habitat (Wilcoxon signed rank test P = 0.007). Within the shared habitat, the two taxa were separated at fine-scale based on oceanographic features.     

A NEW SPECIES OF BEAKED WHALE MESOPLODON PERUVIANUS SP. N. (CETACEA: ZIPHIIDAE) FROM PERU

Mesoplodon peruvianus , a new species of beaked whale, is described on the basis of ten specimens which have either stranded or been captured between 11°12'S and 15°19'S latitude along the coasts of the provinces of Lima and Ica, south central Peru. This is the thirteenth living species of Mesoplodon recognized in the world's oceans. The animals that were examined were uniformly gray above, shading to lighter gray below. This whale is the smallest species of Mesoplodon (maximum body length 3.72 m) and is characterized by its teeth, which are small (31 to 65 mm long), ovate in cross section, and positioned 2.5% to 8.4% of the mandibular length from the anterior extremity, and posterior to the mandibular symphysis.     

Geographic differences in Blainville's beaked whale (Mesoplodon densirostris) echolocation clicks

Aim

Understanding cetacean species' distributions and population structure over space and time is necessary for effective conservation and management. Geographic differences in acoustic signals may provide a line of evidence for population-level discrimination in some cetacean species. We use acoustic recordings collected over broad spatial and temporal scales to investigate whether global variability in echolocation click peak frequency could elucidate population structure in Blainville's beaked whale (Mesoplodon densirostris), a cryptic species well-studied acoustically.

Location

North Pacific, Western North Atlantic and Gulf of Mexico.

Time period

2004–2021.

Major taxa studied

Blainville's beaked whale.

Methods

Passive acoustic data were collected at 76 sites and 150 cumulative years of data were analysed to extract beaked whale echolocation clicks. Using an automated detector and subsequent weighted network clustering on spectral content and interclick interval of clicks, we determined the properties of a primary cluster of clicks with similar characteristics per site. These were compared within regions and across ocean basins and evaluated for suitability as population-level indicators.

Results

Spectral averages obtained from primary clusters of echolocation clicks identified at each site were similar in overall shape but varied in peak frequency by up to 8 kHz. We identified a latitudinal cline, with higher peak frequencies occurring in lower latitudes.

Main conclusions

It may be possible to acoustically delineate populations of Blainville's beaked whales. The documented negative correlation between signal peak frequency and latitude could relate to body size. Body size has been shown to influence signal frequency, with lower frequencies produced by larger animals, which are subsequently more common in higher latitudes for some species, although data are lacking to adequately investigate this for beaked whales. Prey size and depth may shape frequency content of echolocation signals, and larger prey items may occur in higher latitudes, resulting in lower signal frequencies of their predators.     

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.     

A MASS STRANDING OF SPERM WHALES IN OREGON: SEX AND AGE COMPOSITION OF THE SCHOOL

A school of 41 sperm whales, Physeter macrocephalus , that stranded near the mouth of the Siuslaw River, Oregon (43°59'N, 124°08'W), on 16 June 1979 consisted of 13 males and 28 females. Their ages were estimated by assuming that each postnatal dentin layer represents one year. The males were all sub-adults, 9.3–11.5 m long and 14–21 yr old. The females were 9.3–11.4 m long, and 11 to about 58 yr old. Ten females were dissected; nine were sexually mature and three were carrying fetuses 2.64–4.62 m long. None of the 28 females was visibly lactating.     

Molecular studies on two variant repeat types of the common cetacean DNA satellite of the sperm whale, and the relationship between Physeteridae (sperm whales) and Ziphiidae (beaked whales)

In the sperm whale (Physeter macrocephalus) two different repeat types (A and B) of the common cetacean DNA satellite were identified. The evolution of each group of repeats appears to be independent from that of the other. The sequence similarity between the two groups is less than the similarity between group A and repeats of the satellite in related whale species. The systematic relationship within and between the families Physeteridae (sperm whales) and Ziphiidae (beaked whales) was addressed by both sequence analysis of the satellite and comparisons with the families Delphinidae and Phocoenidae. The mysticete blue whale (Balaenoptera musculus) was used as an outgroup in the comparisons. The molecular phylogeny, when maximum-parsimony analysis and the neighbor-joining method were used, grouped together species of each family. At the family level the ziphiids grouped closet to the families Phocoenidae and Delphinidae. The similarities between the common cetacean satellite of the blue whale and the sperm whale were greater than those between the blue whale and the other odontocetes included, suggesting that the evolution of the satellite is slower in the sperm whale than in the other odontocetes.      

Using Energetic Models to Investigate the Survival and Reproduction of Beaked Whales (family Ziphiidae)

Mass stranding of several species of beaked whales (family Ziphiidae) associated with exposure to anthropogenic sounds has raised concern for the conservation of these species. However, little is known about the species’ life histories, prey or habitat requirements. Without this knowledge, it becomes difficult to assess the effects of anthropogenic sound, since there is no way to determine whether the disturbance is impacting the species’ physical or environmental requirements. Here we take a bioenergetics approach to address this gap in our knowledge, as the elusive, deep-diving nature of beaked whales has made it hard to study these effects directly. We develop a model for Ziphiidae linking feeding energetics to the species’ requirements for survival and reproduction, since these life history traits would be the most likely to be impacted by non-lethal disturbances. Our models suggest that beaked whale reproduction requires energy dense prey, and that poor resource availability would lead to an extension of the inter-calving interval. Further, given current information, it seems that some beaked whale species require relatively high quality habitat in order to meet their requirements for survival and reproduction. As a result, even a small non-lethal disturbance that results in displacement of whales from preferred habitats could potentially impact a population if a significant proportion of that population was affected. We explored the impact of varying ecological parameters and model assumptions on survival and reproduction, and find that calf and fetus survival appear more readily affected than the survival of adult females.     

Deep-diving beaked whales dive together but forage apart

Echolocating animals that forage in social groups can potentially benefit from eavesdropping on other group members, cooperative foraging or social defence, but may also face problems of acoustic interference and intra-group competition for prey. Here, we investigate these potential trade-offs of sociality for extreme deep-diving Blainville′s and Cuvier''s beaked whales. These species perform highly synchronous group dives as a presumed predator-avoidance behaviour, but the benefits and costs of this on foraging have not been investigated. We show that group members could hear their companions for a median of at least 91% of the vocal foraging phase of their dives. This enables whales to coordinate their mean travel direction despite differing individual headings as they pursue prey on a minute-by-minute basis. While beaked whales coordinate their echolocation-based foraging periods tightly, individual click and buzz rates are both independent of the number of whales in the group. Thus, their foraging performance is not affected by intra-group competition or interference from group members, and they do not seem to capitalize directly on eavesdropping on the echoes produced by the echolocation clicks of their companions. We conclude that the close diving and vocal synchronization of beaked whale groups that quantitatively reduces predation risk has little impact on foraging performance.     

Acoustic recordings,biological observations,and genetic identification of a rare(?) beaked whale in the North Pacific: Mesoplodon carlhubbsi

Although Hubbs' beaked whale (Mesoplodon carlhubbsi) was previously known from over 60 strandings on both sides of the North Pacific, it had been identified alive in the wild only once, off Oregon in 1994. In September 2021, we conducted a search effort for beaked whales off the coast of Oregon using a towed hydrophone array and a visual search team. Approximately 350 km off the Columbia River mouth, we detected the vocalizations of an unidentified mesoplodont whale; we stopped our vessel and waited in the area until two unidentified juvenile Mesoplodon surfaced and stayed near our vessel for almost 2 hr. During that time, we took numerous photographs and videos, made behavioral observations, and recorded their vocalizations. The DNA sequence from a biopsy sample identified them as M. carlhubbsi. In this paper, we discuss our biological observations, including color patterning and acquired markings, behavioral observations, and describe for the first time the acoustic characteristics of this species. We confirm that M. carlhubbsi is the source of a previously unidentified acoustic signal known as BW37V, and we update what is known about the at-sea distribution of this species based on previous recordings and observational records.     

TWO TYPES OF BLUE WHALE CALLS RECORDED IN THE GULF OF ALASKA

At one time blue whales were found throughout the Gulf of Alaska, however, none have been sighted there in post-whaling era surveys. To determine if blue whales ( Balaenoptera musculus ) might now occur in the Gulf of Alaska, an array of hydrophones was deployed there in October 1999. Data were retrieved in May 2000 and in June 2001. Spectrograms from a random subsample comprising 15% of the ∼63,000 h of data were visually examined for blue whale calls. Call types attributed to both northeastern and northwestern Pacific blue whales were recorded. Both of these call types were recorded seasonally from the initial deployment date in October 1999 through the third week of December 1999 and then from July 2000 through mid-December 2000. Both call types were regularly recorded on the same hydrophone at the same time indicating clear temporal and spatial overlap of the animals producing these calls. Two blue whale call types were recorded in the Gulf of Alaska suggesting that perhaps two stocks use this area. The northeastern call type has now been documented from the equator up to at least 55°N in the eastern North Pacific.     

Vessel noise affects beaked whale behavior: results of a dedicated acoustic response study

Some beaked whale species are susceptible to the detrimental effects of anthropogenic noise. Most studies have concentrated on the effects of military sonar, but other forms of acoustic disturbance (e.g. shipping noise) may disrupt behavior. An experiment involving the exposure of target whale groups to intense vessel-generated noise tested how these exposures influenced the foraging behavior of Blainville's beaked whales (Mesoplodon densirostris) in the Tongue of the Ocean (Bahamas). A military array of bottom-mounted hydrophones was used to measure the response based upon changes in the spatial and temporal pattern of vocalizations. The archived acoustic data were used to compute metrics of the echolocation-based foraging behavior for 16 targeted groups, 10 groups further away on the range, and 26 non-exposed groups. The duration of foraging bouts was not significantly affected by the exposure. Changes in the hydrophone over which the group was most frequently detected occurred as the animals moved around within a foraging bout, and their number was significantly less the closer the whales were to the sound source. Non-exposed groups also had significantly more changes in the primary hydrophone than exposed groups irrespective of distance. Our results suggested that broadband ship noise caused a significant change in beaked whale behavior up to at least 5.2 kilometers away from the vessel. The observed change could potentially correspond to a restriction in the movement of groups, a period of more directional travel, a reduction in the number of individuals clicking within the group, or a response to changes in prey movement.