Humpback whales (Megaptera novaeangliae) attend both Mexico and Hawaii breeding grounds in the same winter: mixing in the northeast Pacific

Humpback whales that assemble on winter breeding grounds in Mexico and Hawaii have been presumed to be, at least, seasonally isolated. Recently, these assemblies were declared Distinct Population Segments under the US Endangered Species Act. We report two humpback whales attending both breeding grounds in the same season—one moving from Hawaii to Mexico and the other from Mexico to Hawaii. The first was photo-identified in Maui, Hawaii on 23 February 2006 and again, after 53 days and 4545 km, on 17 April 2006 in the Revillagigedo Archipelago, Mexico. The second was photo-identified off Guerrero, Mexico on 16 February 2018 and again, 49 days and 5944 km later, on 6 April 2018 off Maui. The 2006 whale was identified in summer off Kodiak Island, Alaska; the 2018 whale off British Columbia. These Mexico–Hawaii identifications provide definitive evidence that whales in these two winter assemblies may mix during one winter season. This, combined with other lines of evidence on Mexico–Hawaii mixing, including interchange of individuals year to year, long-term similarity of everchanging songs, one earlier same-season travel record, and detection of humpback whales mid-ocean between these locations in winter, suggests reassessment of the ‘distinctiveness'' of these populations may be warranted.     

SPATIAL AND SEASONAL DISTRIBUTION OF THE HUMPBACK WHALE, MEGAPTERA NOVAEANGLIAE, IN THE MEXICAN PACIFIC

From observations of the spatial distribution of humpback whales in the Mexican Pacific between 1981 and 1986, it is possible to recognize four subregions: 1) the southern coast of Baja California; 2) the northern Gulf of California, including the Midriff Islands; 3) the mainland coast of Mexico, including the Isla Isabel and Islas Tres Marias and 4) the Revillagigedo Archipelago. The seasonal distribution of whales near the Mexican mainland and the Revillagigedo Archipelago extends from November to May and is similar to that of other winter breeding grounds, including the Hawaiian Islands. Along the southern coast of Baja California, whales have been observed from September to April, possibly indicating a shorter migratory route. In the northern Gulf of California, however, humpback whales have been reported throughout the year and are occasionally observed feeding during both summer and winter months. The degree of individual movement between the four subregions is still unknown. The number of individual humpback whales identified photographically in recent years suggests that there ate more whales in the Mexican Pacific than previously reported.     

Distribution and migratory destinations of humpback whales off the Pacific coast of Central America during the boreal winters of 1996–2003

Here, we examine the distribution, habitat use, and migratory destinations of North Pacific humpback whales wintering off Central America. Coastal boat surveys were conducted off Costa Rica and Panama between 1996 and 2003. In 1999, a broader survey was conducted along most of Central America. Over 23,000 km were surveyed, with the greatest effort off southern Costa Rica. We made 191 sightings of 320 individual humpback whales. Whales were seen between 14°N and 8°N, making this the most southerly of the North Pacific wintering areas. Encounters included singles, adult pairs, singers, and mother/calf pairs. Mother/calf pairs accounted for 27% of all groups sighted, which is one of the highest sighting rates reported among North Pacific wintering areas. Sixty percent of sightings occurred in depths <50 m. Average sea surface temperature was 28.6°C (±1.0 SD). Ninety percent of the 77 unique whales photo‐identified were also seen in the California–Oregon–Washington feeding aggregation. The 1999 survey showed that humpback whales were widely distributed along the Central American coast at relatively low densities. The extensive distribution of animals, the higher proportion of calves, and the almost exclusive migration to a single feeding area contrast with observations in other regions.     

MOVEMENTS OF NORTH PACIFIC BLUE WHALES DURING THE FEEDING SEASON OFF SOUTHERN CALIFORNIA AND THEIR SOUTHERN FALL MIGRATION1

The satellite-acquired locations of 10 blue whales (Balaenoptera musculus) tagged off southern California with Argos radio tags were used to identify (1) their movements during the late summer feeding season; (2) the routes and rate of travel for individuals on their southern fall migration; and (3) a possible winter calving/breeding area. Whales were tracked from 5.1 to 78.1 d and from 393 to 8,668 km. While in the Southern California Bight, most of the locations for individual whales were either clumped or zigzagged in pattern, suggesting feeding or foraging (searching for prey). Average speeds ranged from 2.4 to 7.2 km/h. One whale moved north to Cape Mendocino, and four migrated south along the Baja California, Mexico coast, two passing south of Cabo San Lucas on the same day. One of the latter whales traveled an additional 2,959 km south in 30.5 d to within 450 km of the Costa Rican Dome (CRD), an upwelling feature. The timing of this migration suggests the CRD may be a calving/breeding area for North Pacific blue whales. Although blue whales have previously been sighted in the Eastern Tropical Pacific (ETP), this is the first evidence that whales from the feeding aggregation off California range that far south. The productivity of the CRD may allow blue whales to feed during their winter calving/breeding season, unlike gray whales (Eschrichtius robustus) and humpbacks (Megaptera novaeangliae) which fast during that period.     

HISTORICAL OBSERVATIONS OF HUMPBACK AND BLUE WHALES IN THE NORTH ATLANTIC OCEAN: CLUES TO MIGRATORY ROUTES AND POSSIBLY ADDITIONAL FEEDING GROUNDS

The seasonal distributions of humpback and blue whales ( Megaptera novaeangliae and Balaenoptera musculus , respectively) in the North Atlantic Ocean are not fully understood. Although humpbacks have been studied intensively in nearshore or coastal feeding and breeding areas, their migratory movements between these areas have been largely inferred. Blue whales have only been studied intensively along the north shore of the Gulf of St. Lawrence, and their seasonal occurrence and movements elsewhere in the North Atlantic are poorly known. We investigated the historical seasonal distributions of these two species using sighting and catch data extracted from American 18th and 19th century whaling logbooks. These data suggest that humpback whales migrated seasonally from low-latitude calving/ breeding grounds over a protracted period, and that some of them traveled far offshore rather than following coastal routes. Also, at least some humpbacks apparently fed early in the summer west of the Mid-Atlantic Ridge, well south of their known present-day feeding grounds. In assessing the present status of the North Atlantic humpback population, it will be important to determine whether such offshore feeding does in fact occur. Blue whales were present across the southern half of the North Atlantic during the autumn and winter months, and farther north in spring and summer, but we had too few data points to support inferences about these whales' migratory timing and routes.     

SWIMMING SPEEDS OF SINGING AND NON-SINGING HUMPBACK WHALES DURING MIGRATION

Limited data exist on swimming speeds of humpback whales ( Megaptera novaeangliae ) and none on swimming speeds of singing whales during migration. We tracked humpback whales visually and acoustically during migration from the breeding grounds past our study site on the east coast of Australia (latitude 26°28'S). The mean swimming speed for whales while singing was 2.5 km/h, significantly less than for non-singing whales with a mean of 4.0 km/h but significantly greater than the mean of 1.6 km/h observed for singing whales on the Hawaiian breeding grounds. Between song sessions, there was no significant difference in speeds between whales that had been singing and other whales. Migration speeds were less for whales while singing but increased during the season. Although humpback whales can swim rapidly while singing (maximum observed 15.6 km/h), they generally do not do so, even during migration. Slower migration by singers would delay their return to the polar feeding areas and may be costly, but may be a strategy to provide access to more females.     

Critically endangered western gray whales migrate to the eastern North Pacific

Western North Pacific gray whales (WGWs), once considered extinct, are critically endangered with unknown migratory routes and reproductive areas. We attached satellite-monitored tags to seven WGWs on their primary feeding ground off Sakhalin Island, Russia, three of which subsequently migrated to regions occupied by non-endangered eastern gray whales (EGWs). A female with the longest-lasting tag visited all three major EGW reproductive areas off Baja California, Mexico, before returning to Sakhalin Island the following spring. Her 22 511 km round-trip is the longest documented mammal migration and strongly suggests that some presumed WGWs are actually EGWs foraging in areas historically attributed to WGWs. The observed migration routes provide evidence of navigational skills across open water that break the near-shore north–south migratory paradigm of EGWs. Despite evidence of genetic differentiation, these tagging data indicate that the population identity of whales off Sakhalin Island needs further evaluation.     

Southern Hemisphere humpback whales wintering off Central America: insights from water temperature into the longest mammalian migration

We report on a wintering area off the Pacific coast of Central America for humpback whales (Megaptera novaeangliae) migrating from feeding areas off Antarctica. We document seven individuals, including a mother/calf pair, that made this migration (approx. 8300km), the longest movement undertaken by any mammal. Whales were observed as far north as 11 degrees N off Costa Rica, in an area also used by a boreal population during the opposite winter season, resulting in unique spatial overlap between Northern and Southern Hemisphere populations. The occurrence of such a northerly wintering area is coincident with the development of an equatorial tongue of cold water in the eastern South Pacific, a pattern that is repeated in the eastern South Atlantic. A survey of location and water temperature at the wintering areas worldwide indicates that they are found in warm waters (21.1-28.3 degrees C), irrespective of latitude. We contend that while availability of suitable reproductive habitat in the wintering areas is important at the fine scale, water temperature influences whale distribution at the basin scale. Calf development in warm water may lead to larger adult size and increased reproductive success, a strategy that supports the energy conservation hypothesis as a reason for migration.     

NORTH PACIFIC HUMPBACK WHALE SONGS: A COMPARISON OF SOUTHEAST ALASKAN FEEDING GROUND SONGS WITH HAWAIIAN WINTERING GROUND SONGS

Humpback whales sing long, complex songs on their wintering grounds. On 25 August 1979 and 3 September 1981, we made recordings of humpback whale songs in southeastern Alaska, showing that humpback whales also sing on the summer feeding grounds. Both these Alaskan samples are songs in that they are repeating cyclical sound patterns and follow the known structure for humpback whale song. The Alaskan songs contain all the same material sung in the same order as that heard off Mexico and Hawaii in the surrounding wintering seasons. However, song, theme and some phrase durations are abbreviated in the Alaskan songs. The recording of these two songs represents the full sample of song recorded from 155 days over five years of attempting to record humpback whale song in Alaskan waters.     

SPERM WHALE SURFACE ACTIVITY FROM TRACKING BY RADIO AND SATELLITE TAGS1

Three 12-m sperm whales (Physeter catodon) were tagged and tracked west of Dominica in the southeast Caribbean to follow the surfacing patterns and movements of these presumed subadult males. Whale N was tagged in April 1993 with a 30-MHz radio tag and tracked for two days. Whale H was tagged in April 1995 with a 30-MHz radio tag and tracked for 4.6 d. Whale A was tagged in April 1995 with a satellite-monitored tag tracked by ARGOS for 21.5 d, the first four of which were concurrent with the tracking of Whale H, an associate. The tagged whales remained west of Dominica for at least 2, 5, and 13 d, respectively. Whales N and A then moved southward to waters off Martinique. There were no apparent effects on the whales by tagging or the presence of the tags. The whales averaged speeds of 2.6-3.5 km/h. Surfacings, indicated by tag signals, were of two types: short surfacings apparently primarily for respiration, averaging 7-10.5 min between repeated longer dives, occurring day and night; and extended surfacings seemingly for rest and social interactions with conspecifics, occurring mostly in daylight. Whales were near the surface for 20.4%–22.6% of the total time (26.6%–27.1% during the day and 14.9%–17.1% at night). Delayed blowing was observed as Whale N surfaced for 8.3 min between 47- and 45-min dives but delayed the first of its 31 blows for 1.5 min after surfacing.     

Population structure and accompanying biota of the snail Turbo (Callopoma) funiculosus (Gastropoda: Turbinidae), on Socorro Island, Revillagigedo Archipelago, Mexico

The porcelain snail, Turbo funiculosus, is a potential fishery resource that almost has not been studied or used commercially. In March of 1992, we sampled T. funiculosus in Bahia Binners and Bahia Blanca, Socorro Island, Revillagigedo Archipelago, using 25 m2 quadrants. We found that total length varies between 3.0 and 85 mm, proportional weight between 3.0 and 228 g, and density between 6.21 and 9.87 ind/m2. The largest organisms (over 50 mm) contain 30.6% of soft parts. Porcelain snail populations remain unexploited on Revillagedo Archipelago and could be a potential resource under an adequate management strategy.     

MOVEMENTS AND POPULATION STRUCTURE OF HUMPBACK WHALES IN THE NORTH PACIFIC

Despite the extensive use of photographic identification methods to investigate humpback whales in the North Pacific, few quantitative analyses have been conducted. We report on a comprehensive analysis of interchange in the North Pacific among three wintering regions (Mexico, Hawaii, and Japan) each with two to three subareas, and feeding areas that extended from southern California to the Aleutian Islands. Of the 6,413 identification photographs of humpback whales obtained by 16 independent research groups between 1990 and 1993 and examined for this study, 3,650 photographs were determined to be of suitable quality. A total of 1,241 matches was found by two independent matching teams, identifying 2,712 unique whales in the sample (seen one to five times). Site fidelity was greatest at feeding areas where there was a high rate of resightings in the same area in different years and a low rate of interchange among different areas. Migrations between winter regions and feeding areas did not follow a simple pattern, although highest match rates were found for whales that moved between Hawaii and southeastern Alaska, and between mainland and Baja Mexico and California. Interchange among subareas of the three primary wintering regions was extensive for Hawaii, variable (depending on subareas) for Mexico, and low for Japan and reflected the relative distances among subareas. Interchange among these primary wintering regions was rare. This study provides the first quantitative assessment of the migratory structure of humpback whales in the entire North Pacific basin.     

Migratory preferences of humpback whales between feeding and breeding grounds in the eastern South Pacific

Latitudinal preferences within the breeding range have been suggested for Breeding Stock G humpback whales that summer in different feeding areas of the eastern South Pacific. To address this hypothesis, humpback whales photo‐identified from the Antarctic Peninsula and the Fueguian Archipelago (southern Chile) were compared with whales photo‐identified from lower latitudes extending from northern Peru to Costa Rica. This comparison was performed over a time span that includes 18 austral seasons. A total of 238 whales identified from the Antarctic Peninsula and 25 whales from the Fueguian Archipelago were among those photo‐identified at the breeding grounds. Our findings showed that humpback whales from each feeding area were resighted unevenly across the breeding grounds, which suggests a degree of spatial structuring in the migratory pathway. Humpback whales that feed at the Antarctic Peninsula were more likely to migrate to the southern breeding range between northern Peru and Colombia, whereas whales that feed at the Fueguian Archipelago were more likely to be found in the northern range of the breeding ground off Panama. Further photo‐identification efforts and genetic sampling from poorly sampled or unsampled areas are recommended to confirm these reported connectivity patterns.     

Changes in North Atlantic right whale (Eubalaena glacialis) cow–calf association times and use of the calving ground: 1993–2005

Like most mysticetes, North Atlantic right whale cows generally separate from their calves on their feeding grounds within a year. Right whale life history data from 1993 to 2005 were analyzed to determine the duration of cow/calf associations and where the pair separated. A change occurred with the 2001 cows; 71% of those available stayed with their calves into the second year and this behavior remained elevated for several years. Less experienced cows, independent of their age, were more likely to extend their associations. The occurrence of cow/yearling associations was not related to the length of the cow's previous interbirth interval, used as a proxy for cow condition, but the hypothesis that body condition impacts how long cows nurse their young could not be adequately tested. Seventy-seven percent of the observed cow/yearling pairs also returned to the calving ground, a substantial physiological investment given the 1,450 km plus migration and the fact that they fast there, indicating that factors other than nutrition also influenced the cow's behavior. The concurrent increase in juveniles in the shallow waters of the winter calving grounds may afford naive whales greater protection from predators or provide a social benefit that improves their overall fitness.     

The effect of wind, season and latitude on the migration speed of white storks Ciconia ciconia, along the eastern migration route

The relation between wind, latitude and daily migration speed along the entire migration route of white storks was analysed. Mean daily migration speed was calculated using satellite telemetry data for autumn and spring migration of white storks from their breeding grounds in Germany and Poland to wintering grounds in Africa and back. The National Center for Environmental Prediction (NCEP) reanalysis data were used to systematically fit 850 mb wind vectors to daily migration speed along the migration route. White storks migrated significantly faster and had a shorter migration season in autumn (10 km/h) compared to spring (6.4 km/h). In autumn mean daily migration speed was significantly slower in Europe (8.0 km/h) than in the Middle East (11.1 km/h) and Africa (11.0 km/h). In spring mean daily migration speed was significantly faster in Africa (10.5 km/h) as birds left their wintering grounds than in the Middle East (4.3 km/h). Migration speed then increased in Europe (6.5 km/h) as birds approached their breeding grounds. In both spring and autumn tailwind (at 850mb) and latitude were found to be significant variables related to daily migration speed.     

DIEL VARIATION IN MIGRATION RATES OF EASTERN PACIFIC GRAY WHALES MEASURED WITH THERMAL IMAGING SENSORS

We recorded the blows of gray whales during their southbound migration past central California in January 1994, 1995, and 1996, using thermal imaging sensors. For our sampling purposes, we defined day (0730–1630) and night (1630–0730) to coincide with the on/off effort periods of the visual counts being conducted concurrently. We pooled data across the three years of sampling and tested for diel variation in surfacing interval, pod size, offshore distance, migration rate, and swimming speed by comparing paired day/night means for samples collected within the respective 24-h period. We performed these tests using data from the entire migration period and then repeated the tests for samples collected prior to and after the approximate median migration date (15 January). Over the entire migration period we observed larger diurnal pod sizes (x?_day= 1.75 ± 0.280, x?_night= 1.63 ± 0.232) and greater diurnal offshore distances (x?_day= 2.30 ± 0.328 km, x?_night= 2.03 ± 0.356 km) but found no diel variation in surfacing interval. For the entire migration period, the nocturnal migration rate (average number of whales passing per hour) was higher than the diurnal rate. During the first half of the migration we detected no diel variation in pod size or surfacing interval, but diurnal offshore distances were larger than at night (x?_day= 2.28 ± 0.273 km, x?_night= 1.96 ± 0.318 km). Diurnal and nocturnal migration rates prior to 15 January were not different. During the second half of the migration, there was no diel variation in surfacing interval, pod size, or distance offshore, but the nocturnal migration rate was higher (28%, SE = 11.6%) than the diurnal rate. We found no diel variation in swimming speed in any comparison. We propose that later migrants socialize more during the day, which effectively slows their diurnal rate of migration relative to nocturnal rates.     

Population structure of nuclear and mitochondrial DNA variation among humpback whales in the North Pacific

The population structure of variation in a nuclear actin intron and the control region of mitochondrial DNA is described for humpback whales from eight regions in the North Pacific Ocean: central California, Baja Peninsula, nearshore Mexico (Bahia Banderas), offshore Mexico (Socorro Island), southeastern Alaska, central Alaska (Prince Williams Sound), Hawaii and Japan (Ogasawara Islands). Primary mtDNA haplotypes and intron alleles were identified using selected restriction fragment length polymorphisms of target sequences amplified by the polymerase chain reaction (PCR–RFLP). There was little evidence of heterogeneity in the frequencies of mtDNA haplotypes or actin intron alleles due to the year or sex composition of the sample. However, frequencies of four mtDNA haplotypes showed marked regional differences in their distributions (Φ_ST = 0.277; P < 0.001; n = 205 individuals) while the two alleles showed significant, but less marked, regional differences (Φ_ST = 0.033; P < 0.013; n = 400 chromosomes). An hierarchical analysis of variance in frequencies of haplotypes and alleles supported the grouping of six regions into a central and eastern stock with further partitioning of variance among regions within stocks for haplotypes but not for alleles. Based on available genetic and demographic evidence, the southeastern Alaska and central California feeding grounds were selected for additional analyses of nuclear differentiation using allelic variation at four microsatellite loci. All four loci showed significant differences in allele frequencies (overall F_ST = 0.043; P < 0.001; average n = 139 chromosomes per locus), indicating at least partial reproductive isolation between the two regions as well as the segregation of mtDNA lineages. Although the two feeding grounds were not panmictic for nuclear or mitochondrial loci, estimates of long-term migration rates suggested that male-mediated gene flow was several-fold greater than female gene flow. These results include and extend the range and sample size of previously published work, providing additional evidence for the significance of genetic management units within oceanic populations of humpback whales.     

Mother Knows Best: Occurrence and Associations of Resighted Humpback Whales Suggest Maternally Derived Fidelity to a Southern Hemisphere Coastal Feeding Ground

Site fidelity is common among migratory cetaceans, including humpback whales (Megaptera novaeangliae). In the Northern Hemisphere it has been found that fidelity to humpback whale feeding grounds is transferred maternally but this has never been shown for the species in the Southern Hemisphere. We examined this in a unique feeding area off west South Africa using resighting data of 68 individually identified humpback whales by means of photographic (tail flukes and dorsal fins) and/or molecular methods (microsatellite genotyping) over an 18 year span. We found short-term association patterns and recurrent visits typical of other feeding grounds. Males and females had different seasonality of attendance. Significant female-dominated presence corresponded to timing of an expected influx of females on their southward migration from the breeding ground: firstly non-nursing (possibly pregnant) females in mid-spring, and mothers and calves in mid-to late summer. The potential benefit of this mid-latitude feeding area for females is illustrated by a record of a cow with known age of at least 23 years that produced calves in three consecutive years, each of which survived to at least six months of age: the first record of successful post-partum ovulation for this species in the Southern Hemisphere. We recorded association of a weaned calf with its mother, and a recurring association between a non-lactating female and male over more than two years. Moreover, three animals first identified as calves returned to the same area in subsequent years, sometimes on the same day as their mothers. This, together with numerous Parent-Offspring relations detected genetically among and between resighted and non-resighted whales is strongly suggestive of maternally derived site fidelity at a small spatial scale by a small sub-population of humpback whales.     

Within‐year movements and site fidelity of Purple Sandpipers during the nonbreeding season

ABSTRACT Although within‐year site fidelity to specific wintering sites allows shorebirds to use prior knowledge of resources and microhabitats, such fidelity may also make populations more vulnerable to extirpation in the event of increased predation pressure, habitat loss, or disturbance. In the eastern Atlantic, Purple Sandpipers (Calidris maritima) have been found to be highly faithful to specific sites in wintering areas. However, little is known about the use of wintering areas by these sandpipers along the coast of Maine. We quantified movements of 60 radio‐marked Purple Sandpipers in a bay near the mainland and on an offshore cluster of islands along the mid‐coast of Maine during two winters (2005–2006 and 2006–2007). Birds marked in early‐ and mid‐December remained until spring migration, with no evidence of onward migration. Mean maximum distances moved did not differ significantly between either males (8.6 ± 1.0 [SE] km; N= 30) and females (7.4 ± 0.8 km; N= 30) or juveniles (9.9 ± 1.6 km; N= 9) and adults (7.8 ± 1.1 km; N= 26). We also detected no monthly (January–May) differences in maximum distances moved. Sixty percent of marked individuals moved ≤5 km between the two most distant relocations and no birds moved >25 km during the 2‐ to 4‐month tracking period. We attribute the high site fidelity primarily to the plentiful prey base in the study area. During a 2‐d period with severe cold, feeding areas at locations protected from wave action became encased in ice and birds at these locations moved up to 10 km offshore to sites with less ice. Species with strong site fidelity, like wintering Purple Sandpipers, may be at higher risk in the event of large‐scale changes in their food base, increased predation pressure, habitat loss, or disturbance. However, the short‐distance movements made when intertidal feeding areas became encased in ice suggest that Purple Sandpipers could potentially move greater distances in response to changing conditions in their wintering areas.     

Humpback whales interfering when mammal‐eating killer whales attack other species: Mobbing behavior and interspecific altruism?

Humpback whales (Megaptera novaeangliae) are known to interfere with attacking killer whales (Orcinus orca). To investigate why, we reviewed accounts of 115 interactions between them. Humpbacks initiated the majority of interactions (57% vs. 43%; n = 72), although the killer whales were almost exclusively mammal‐eating forms (MEKWs, 95%) vs. fish‐eaters (5%; n = 108). When MEKWs approached humpbacks (n = 27), they attacked 85% of the time and targeted only calves. When humpbacks approached killer whales (n = 41), 93% were MEKWs, and ≥87% of them were attacking or feeding on prey at the time. When humpbacks interacted with attacking MEKWs, 11% of the prey were humpbacks and 89% comprised 10 other species, including three cetaceans, six pinnipeds, and one teleost fish. Approaching humpbacks often harassed attacking MEKWs (≥55% of 56 interactions), regardless of the prey species, which we argue was mobbing behavior. Humpback mobbing sometimes allowed MEKW prey, including nonhumpbacks, to escape. We suggest that humpbacks initially responded to vocalizations of attacking MEKWs without knowing the prey species targeted. Although reciprocity or kin selection might explain communal defense of conspecific calves, there was no apparent benefit to humpbacks continuing to interfere when other species were being attacked. Interspecific altruism, even if unintentional, could not be ruled out.