The causes of the stranding of large numbers of whales and dolphins on beaches, resulting in the deaths of most if not all individuals involved, have been an unanswered question at least since the days of Greek philosopher Aristotle (who noted this curious phenomenon in his Historia Animalia in 350 BCE). Various suggestions have been made to explain this, including possible disease, parasitic infection, sympathetic mass suicide, electromagnetic disorientation, and anthropogenic activity (Cordes, 1982, Sunduram et al., 2006). It is clear that in the great majority of mass-strandings, many of the animals that arrive together on a beach are not near death, as they appear to swim away competently once moved to deeper water by rescuers (though some may return to the beach). Many species mass-strand, but with rare exceptions (e.g. D’Amico et al. 2009) each of the mass stranding events involves only one species, and the numbers that strand together appear to reflect the size of the social groups in which these animals move around in the oceans. In the case of Long-finned Pilot Whales (Globicephalus melas), the strandings often involve larger numbers than those recorded at sea, but they appear to divide into widely-spaced smaller groups while foraging (Visser et al. 2017), so that average group sizes at sea may be underestimated.
Because cetaceans navigate by sound, one suggested cause has been the possible dysfunction of echo-location, due to the lack of sonic reflection from a gently sloping beach, or because of the absorption of sound by the presence of sand or microbubbles in shallow water (Dudok van Heel, 1962, Chambers and James 2005, Sundaram et al., 2006). These suggestions have yet to be thoroughly tested, but one might expect that animals approaching a beach with a gentle slope would have some warning because they could easily detect that the depth beneath them is small and decreasing.
Suggestions that geomagnetic topography may be having an effect can be discounted, at least in the New Zealand environment, as herd strandings have been shown to have no relationship to geomagnetic contours or magnetic minima (Brabyn 1991, Brabyn & Frew, 1994). The relation to geomagnetic storms is also rather weak for New Zealand (Pulkinnen et al. 2020, Vanselow 2020).
A simple explanation, one that appears most likely to us, is that one or more individuals become sick (incompetent) and drift onto the shore, and group members follow these incapacitated individuals as they drift onto the shore, due to social bonds that are not yet fully understood. Reggente et al. (2016) have described such nurturing behaviour, and Brabyn (1991) and Brabyn & McLean (1992) describe observations from data prior to mid 1989, which are indicative of the results presented here. Brabyn (1991), in particular, noted that offshore species are more prone to herd-strand than inshore species, and gave a description of these phenomena up to that date. We have gathered evidence for this hypothesis using massed stranding events in New Zealand over the past 40 years.
The New Zealand data set
Stranding events have been recorded in New Zealand dating back to the1840’s, and the record-keeping has been a legal requirement since 1978. They cover a large number of the species that have been recorded to strand both individually and in large groups. Brabyn & McLean (1992), and more recently Betty et al. (2020), have provided detailed descriptions of the spatial and temporal distribution of these stranding events, identifying “hot spots” and temporal records, and focussing on the Long-finned Pilot Whale (LFPW), which is by far the most significant species for strandings in large numbers in New Zealand waters. The database used here was provided by the New Zealand Department of Conservation, via Mr. Mike Ogle, Hannah Hendricks and Project Jonah (projectjonah.org.nz). The data contain some 3782 stranding events, dating from 1980 to the end of 2019. The great majority of these recorded events involve single animals, spread over the whole range of species in New Zealand waters, as described by Betty et al. (2020).
While mass strandings are often defined as involving 3 or more animals, we focus our analysis on events that involve 10 or more stranded animals, dated from 1980 onwards, as such numbers are unlikely to involve chance simultaneous strandings of individual animals or mother-calf pairs, and wind and wave data were available from 1980. This gives a data set of 125 massed stranding events, each involving one species, with numbers of animals in each event ranging from 10 to 616. All mass strandings appear to involve both young and older individuals. During the 1980-2019 period there were 24 events with 100 or more stranded animals, with the largest number of stranded animals in a single event being 616 in the Farewell Spit region. All of these large events involved pilot whales.
The social cultures of cetaceans
Species of whales and dolphins vary in their social structures, many of which are not fully understood. Pilot whales represent the majority of the massed strandings considered here, so some details of their life-style are relevant. These animals are born in pods that may number up to several hundred animals. Long-fin Pilot Whales found in New Zealand take 11 to 15 years to mature to adulthood (Betty et al. 2019). In the North Atlantic sub-species, both the males and females in the pod are born in the pod, but none of the male members are fathers (Amos et al., 1991, 1993). As the culture of the Globicephala melas subspecies appear to be very similar, we assume that the same applies for the South Pacific subspecies. Males must leave the pod to breed with females in other pods. Females born in the pod remain there, and males may or may not return to their home pod.
Members of the pod, most prominently females, generally help younger pod members, even though they may not be the parent (Amos et al.,1991; Oremus et al., 2013; Augusto et al., 2017). Thus although Oremus et al. (2013) have shown that mass strandings in New Zealand and Tasmania involve multiple matrilineal lineages, and mothers and calves are not stranded close to each other, large groups of pilot whales might well stay with a dying senior pod member due to a strong cultural relationship between young and older pod members.
Species other than pilot whales that mass strand in New Zealand may also live for decades in pods, so that it seems possible that when senior members eventually become incapacitated, if they have been prominent in pod social structure, younger pod members might accompany the sick or dying pod member as it drifts, due to social bonds. The cetacean subfamily Globicephalinae includes killer whales, false killer whales and pygmy killer whales as well as the pilot whales. Each of these species mass strands and exhibits strong social structure (Martien et al. 2017). Kinship groups also occur in sperm whales (Richard et al. 1996). In some mass stranding species, the groups that swim together are not all closely related individuals, so that kin selection alone does not explain the social bonds (Ball et al., 2017; Martien et al., 2014, Kobayashi et al.,2020; Patel et al., 2017; Westbury et al., 2021). But Norris and Schilt (1988) and Moller (2012) discuss how cooperative and altruistic patterns in cetaceans could evolve to extend beyond familial units.
It is indisputable that groups of competent individuals of these species do become stranded, and we are interested in determining whether this may be because they accompany one or more dying and bloated individuals. We note the increasing number of impressive studies of the genetic and the social structure of cetacean species, and this field is still developing (Moller, 2012, Reeves et al. 2022).
For all cetacean species, when the animals die it is not uncommon for their floating cadavers to be deposited on nearby coasts. This is reflected by the large number of single strandings, over all relevant species in the New Zealand records. Over the 40-year period considered, there are 3063 recorded single strandings, including 180 cases of single stranded pilot whales.
The next most common stranding number is two animals. In contrast with some other species, stranding events consisting of two pilot whales alone are relatively rare – over the 40-year period there are 8 recorded stranded-pairs of pilot whales out of a total of 196. This is consistent with evidence for the northern sub-species that they tend not to form strong pair-bond relationships (as other species may do), but the males (in particular) can exist as individuals outside a pod, or have a close relationship within a large pod community (Amos et al., 1993). Nor are pilot whales over-represented in stranded groups containing 3 to 9 animals. In a total of 115 stranding events in this range over 40 years, 16 of these events involve pilot whales, which is a number comparable with those of other species.
The stranding process- Stage 1: wind and waves on a floating body
We have examined the fit of the data (particularly the pilot whale data) to the hypothesis that the massed strandings are due to the beaching of one or more pod members who have become ill (for whatever reason), and whose body has become buoyant (for example due to bloating of the gut), and floats on the surface. The motion of such a floating object will be subject to the effects of both the wind and waves at the surface. If a coastline is nearby, the direction of the wind and waves will largely determine whether the floating animal is driven towards the shoreline.
In deep water, where waves are present but not breaking, the main effect of waves on the mean motion of floating bodies is via Stokes drift. This is a mean motion of particles on the surface of the water due to the nonlinear dynamics of sufficiently large waves, which is given by (Craik, 2005)
Here us is the resultant mean speed of the floating object/body in the direction of the waves, α is the wave amplitude, λ the wavelength and T the wave period. Realistically, this gives speeds of magnitude 2 cm/s, which is approaching 2 km/day.
The above gives mean surface speeds due to non-breaking waves, but if the waves are breaking, the speed of the floating object will be much larger. This would be the case when the object reaches the wave surf zone.
The effect of the wind on floating object movement can be estimated as the drag, and this can be somewhat larger than the wave effect in the deep sea. To some extent these two factors (wind and waves) tend to be aligned, as the wind generates the waves. The wind direction can change more quickly, but there is a tendency for the two factors to be aligned, given sufficient steady wind. This consideration of the effects of wind and waves suggests that floating animals may be driven ashore from nearby seas over a period of about a day, but strandings may be noticed only after some time, so that the wind and waves over the two days prior to a stranding must be considered.
The stranding process- stage 2: pod members accompany the floating animal
Our hypothesis assumes that pod members would accompany a senior pod member who is incapacitated and floating. This would depend on the existence of strong social bonds in the group, and we note that many authors have described such social bonds, due to kinship in various cetaceansin the sub-family Globicephalinae (Augusto et al., 2017, Martien et al. 2017), or more complex relations in sperm whales (Whitehead 1996, Richard et al. 1996) and dolphins (Moller 2012) but we have no direct evidence that such bonds result in pod members of each species following incapacitated animals.
The stranding process- stage 3: the beach slope and the tides
Where the coastline consists of steep cliffs or rocky environments, the floating body may remain in the water, or be thrown up by heavy wave action, and the associated pod would presumably keep it’s distance, and there would be no subsequent record of a mass stranding event. But if it is a gently sloping beach, the body of the injured animal would be gradually pushed into very shallow water.
This third part of the massed stranding process also concerns the tides. If the tidal range is significant, and the tide is fairly high as a dead or incapacitated animal arrives, it will be driven to a high level on the beach. If the tide is ebbing, both the dying individual and the accompanying pod members will rapidly become stranded on the beach. The offshore species such as pilot whales will have no significant familiarity with tides, and hence not be aware that the sea level may change by as much as several metres. On a gently sloping beach, a tidal range of even a metre may cause the local shoreline to retreat laterally by several metres in a matter of minutes.
In summary, our hypothesis for the massed stranding phenomenon is a three-stage process: (1) that the wind and waves in the days before the stranding would drive a floating body ashore at the stranding site; (2) that the dying or incapacitated animal would be accompanied closely by many others from the group associated with it; and (3) that the beach at the time when the body arrives would be gently sloping, with the tide at least moderately high at the time, and the tide range sufficient so that the water level retreats quickly.