Month: February 2020

Makah Gray Whale Hunt Waiver – a long-time coming, but still premature?

By Lisa Hildebrand, MSc student, OSU Department of Fisheries & Wildlife, Marine Mammal Institute, Geospatial Ecology of Marine Megafauna Lab

Archaeological site of Ozette Village. Source: Makah Museum.

The Makah, an indigenous people of the Pacific Northwest Coast living in Washington State, have a long history with whaling. Deposits from a mudslide in the village of Ozette suggest that whaling may date back 2,000 years as archaeologists uncovered humpback and gray whale bones and barbs from harpoons (Kirk 1986). However, the history of Makah whaling is also quite recent. On January 29 of this year, the National Marine Fisheries Service (NMFS; informally known as NOAA Fisheries) announced a 45-day public comment period regarding a NMFS proposed waiver on the Marine Mammal Protection Act’s (MMPA) moratorium on the take of marine mammals to allow the Makah to take a limited number of eastern North Pacific gray whales (ENP). To understand how the process reached this point, we first must go back to 1855.

1855 marks the year in which the U.S. government and the Makah entered into the Treaty of Neah Bay (in Washington state). The Makah ceded thousands of acres of land to the U.S. government, and in return reserved their right to whale. Following the treaty, the Makah hunt of gray whales continued until the 1920s. At this point, commercial hunting had greatly reduced the ENP population, so much so that the Makah voluntarily ceased their whaling. The next seven decades brought about the formation of the International Whaling Commission (IWC), the enactment of the Whaling Convention Act, the listing of gray whales as endangered under the U.S. Endangered Species Act, and the enactment of the MMPA. For gray whales, these national and international measures were hugely successful, leading to the removal of the ENP from the Federal List of Endangered Wildlife in 1994 when it was determined that the population had recovered to near its estimated original population size.

One year later on May 5, 1995 (just one month after I was born!), the Makah asked the U.S. Department of Commerce to represent its interest to obtain a quota for gray whales from the IWC in order to resume their treaty right for ceremonial and subsistence harvest of the ENP. The U.S. government pursued this request at the next IWC meeting, and subsequently NMFS issued a final Environmental Assessment that found no significant impact to the ENP population if the hunt recommenced. The IWC set a catch limit and NMFS granted the Makah a quota in 1998. In 1999 the Makah hunted, struck and landed an ENP gray whale.

“Makahs cutting up whale, Neah Bay, ca. 1930. Photo by Asahel Curtis, Courtesy UW Special Collections (CUR767)”. Source and caption: History Link.

I will not go into detail about what happened between 1999 and now because frankly, a lot happened, particularly a lot of legal events including summary judgements, appeals, and a lot of other legal jargon that I do not quite understand. If you want to know the specifics of what happened in those two decades, I suggest you look at NMFS’ chronology of the Makah Tribal Whale Hunt. In short, cases brought against NMFS argued that they did not take a “hard [enough] look” at the National Environmental Policy Act when deciding that the Makah could resume the hunt. Consequently, the hunt was put on hold. Yet, in 2005 NMFS received a waiver request from the Makah on the MMPA’s take moratorium and NMFS published a notice of intent to review this request. A lot more happened between that event and now, including on January 29 of this year when NMFS announced the availability of transcripts from the Administrative Law Judge’s (ALJ) hearing (which happened from November 14-21, 2019) on the proposed regulations and waiver to allow the Makah to resume hunting the ENP. We are currently in the middle of the aforementioned 45-day public comment period on the formal rulemaking record. 

It has been 15 years since the Makah requested the waiver and while the decision has not yet been reached, we are likely nearing the end of this long process. This blog has turned into somewhat of a history lesson (not really my intention) but I feel it is important to understand the lengthy and complex history associated with the decision that is probably going to happen sometime this year. My actual intent for this blog is to ruminate on a few questions, some of which remain unanswered in my opinion, that are large and broad, and important to consider. Some of these questions point out gaps in our ecological knowledge regarding gray whales that I believe should be addressed for a truly informed decision to be made on NMFS’ proposed waiver now or anytime in the near future. 

1. Should the Pacific Coast Feeding Group (PCFG) of gray whales be recognized as its own stock?

Currently, the PCFG are considered a part of the ENP stock. This decision was published following a workshop held by a NMFS task force (Weller et al. 2013). The report concluded that based on photo-identification, genetics, tagging, and other data, there was a substantial level of uncertainty in the strength of the evidence to support the independence of the PCFG from the ENP. Nevertheless, mitochondrial genetic data have indicated a differentiation between the PCFG and the ENP, and the exchange rate between the two groups may be small enough for the two to be considered demographically independent (Frasier et al. 2011). Based on all currently available data, it seems that matrilineal fidelity plays a role in creating population structure within and between the PCFG and the ENP, however there has not been any evidence to suggest that whales from one feeding area (i.e. the PCFG range) are reproductively isolated from whales that utilize other feeding areas (i.e. the Arctic ENP feeding grounds) (Lang et al. 2011). Several PCFG researchers do argue that there needs to be recognition of the PCFG as an independent stock. It is clear that more research, especially efforts to link genetic and photo-identification data within and between groups, is required.

ENP gray whales foraging off the coast of Alaska on their main foraging grounds in the Bering Sea. Photo taken by ASAMM/AFSC. Funded by BOEM IAA No. M11PG00033. Source: NMFS.

2. Is emigration/immigration driving PCFG population growth, or is it births/deaths?

It is unclear whether the current PCFG population growth is a consequence of births and deaths that occur within the group (internal dynamics) or whether it is due to immigration and emigration (external dynamics). Likely, it is a combination of the two, however which of the two has more of an effect or is more prevalent? This question is important to answer because if population growth is driven more by external dynamics, then potential losses to the PCFG population due to the Makah hunt may not be as detrimental to the group as a whole. However, if internal dynamics play a bigger role, then the loss of just a few females could have long-term ramifications for the PCFG (Schubert 2019). NMFS has taken precautions to try and avoid such effects. In their proposed waiver, of the cumulative limit of 16 strikes of PCFG whales over the 10-year waiver period, no more than 8 of the strikes may be PCFG females (Yates 2019a). While a great step, it still begs the question how the loss of 8 females, admittedly over a rather long period of time, may affect population dynamics since we do not know what ultimately drives recruitment. Especially when taken together with potential non-lethal effects on whales (further discussed in question 5 below).

“Scarlet” is a PCFG female who has had multiple calves in the decades that researchers have seen her in the PCFG range. Image captured under NOAA/NMFS permit #21678. Source: L Hildebrand.

3. How important are individual patterns within the PCFG, and how might the loss of these individuals affect the population? 

The hunt will be restricted to the Makah Usual & Accustomed fishing area (U&A), which is off the Washington coast. It has been shown that site fidelity among PCFG individuals is strong. In fact, based on the 143 PCFG gray whales observed in nine or more years from 1996 to 2015, 94.4% were seen in at least one of nine different PCFG regions during six or more of the years they were seen (Calambokidis et al. 2017). While high site-fidelity seems to be common for some PCFG individuals in certain regions, interestingly, an analysis of sighting histories of all individuals that utilized the Makah U&A from 1985-2011 revealed that most PCFG whales do not have strong site fidelity to the Makah U&A (Scordino et al. 2017). Only about 20% of the whales were observed in six or more years of the total 26 years of data analyzed. Since high individual site fidelity does not appear to be strong in this area, perhaps a loss of genetic diversity, cultural knowledge, and behavioral individualism is not of great concern.

“Buttons” seems to have a preference for the southern Oregon coast as in the last 5 years the GEMM Lab has conducted research, he has only been sighted in 1 year in Newport but in all 5 years in Port Orford. However, perhaps such preferences are not common among all PCFG whales. Source: F. Sullivan.

4. How has the current UME affected the situation?

The ENP has experienced two Unusual Mortality Events (UMEs) in the past 20 years; one from 1999-2000 and the second began in May 2019. Many questions arise when thinking about the Makah hunt in light of the UME. 

  • What impacts will the current UME have on ENP and PCFG birth rates in subsequent years? 
  • Could the UME lead to shifts in feeding behavior of ENP whales and result in greater use of PCFG range by more individuals?
  • What caused the UME? Shifting prey availability and a changing climate? Or has the ENP reached carrying capacity? 
  • Will UMEs become more frequent in the future with continued warming of the Arctic? 
  • What is the added impact of such periodic UMEs on population trends?
“A gray whale found dead off Point Reyes National Seashore in northern California [during the 2019 UME]. Photo by M. Flannery, California Academy of Sciences.” Source and caption: NMFS.

A key assumption of the model developed by NMFS (Moore 2019) to forecast PCFG population size for the period 2016-2028, is that the population processes underlying the data from 2002-2015 (population size estimates developed by Calambokidis et al. 2017) will be the same during the forecasted period. In other words, it is assuming that PCFG gray whales will experience similar environmental conditions (with similar variation) during the next decade as the previous one, and that there will be no catastrophic events that could drastically affect population dynamics. The UME that is still ongoing could arguably affect population dynamics enough such that they are drastically different to effects on the population dynamics during the previous decade. The cause of  the 1999/2000 UME remains undetermined and the results of the investigation of the current UME will possibly not be available for several years (Yates 2019b). Even though the ENP did rebound following the 1999/2000 UME and the abundance of the PCFG increased during and subsequent to that UME, much has changed in the 20 years since then. Increased noise due to increased vessel traffic and other anthropogenic activities (seismic surveys, pile driving, construction to name a few) as well as increased coastal recreational and commercial fishing, have all contributed to a very different oceanscape than the ENP and PCFG encountered 20 years ago. Furthermore, the climate has changed considerably since then too, which likely has caused changes in the spatial distribution of habitat and quantity, quality, and predictability of prey. All of these factors make it difficult to predict what impact the UME will have now. If such events were to become more frequent in the future or the impacts of such events are greater than anticipated, then the PCFG population forecasts will not have accounted for this change. 

5. What impacts will the hunt and associated training exercises have on energy and stress levels of whales?

The proposed waiver would allow hunts to occur in the following manner: in even-years, the hunting period is from December 1 of an odd-numbered year through May 31 of the following even-numbered year. While in odd-years, the hunt is limited from July to October.

In the even-years, the hunt coincides with the northbound migration toward the foraging grounds for ENP whales and with the arrival of PCFG whales to their foraging grounds near the Makah U&A. During the northbound migration, gray whales are at their most nutritionally stressed state as they have been fasting for several months. They are therefore most vulnerable to energy losses due to disturbance at this point (Villegas-Amtmann 2019). Attempted strikes and training exercises would certainly cause some level of disturbance and stress to the whales. Furthermore, the timing of even-year hunts, means that hunters would likely encounter pregnant females, as they are the first to arrive at foraging grounds. A loss of just ~4% of a pregnant female’s energy budget could cause them to abort the fetus or not produce a calf that year (Villegas-Amtmann 2019).

In odd-years, the Makah hunt will most certainly target PCFG whales as the Makah U&A forms one of the nine PCFG regions where PCFG individuals will be feeding during those months. However, NMFS’ waiver limits the number of strikes during odd-years to 2 (Yates 2019a), which certainly protects the PCFG population.

Stress is a difficult response to quantify in baleen whales and research on stress through hormone analysis is still relatively novel. It is unlikely that a single boat training approach of a gray whale will have an adverse effect on the individual. However, a whale is never just experiencing one disturbance at a time. There are typically many confounding factors that influence a whale’s state. In an ideal world, we would know what all of these factors are and how to recognize these effects. Yet, this is virtually impossible. Therefore, while precautions will be taken to try to minimize harm and stress to the gray whales, there may very well still be unanticipated impacts that we cannot anticipate. 

Gray whale fluke. Image captured under NOAA/NMFS permit #21678. Photo: L Hildebrand.

Final thoughts

Many unknowns still remain about the ENP and PCFG gray whale populations. During the ALJ hearing, both sides tried to deal with these unknowns. After reading testimony from both sides, it is clear to me that some of the unknowns still have not been reconciled. Ultimately, a lot of the questions circle back to the first one I posed above: Are the PCFG an independent stock? If there is independent population structure, then the proposed waiver put forth by NMFS would likely change. While NMFS has certainly taken the PCFG into account during the declarations of several experts at the ALJ hearing and has aired on the side of caution, the fact that the PCFG is considered part of the ENP might underestimate the impact that a resumption of the Makah hunt may have on the PCFG. As you can see, there are still many questions that should be addressed to make fully informed decisions on such an important ruling. While this research may take several years to obtain results, the data are within reach through synthesis and collaboration that will fill these critical knowledge gaps. 

Literature cited

Calambokidis, J. C., J. Laake, and A. Pérez. 2017. Updated analysis of abundance and population structure of seasonal gray whales in the Pacific Northwest, 1996-2015. International Whaling Commission SC/A17/GW/05.

Frasier, T. R., S. M. Koroscil, B. N. White, and J. D. Darling. 2011. Assessment of population substructure in relation to summer feeding ground use in eastern North Pacific gray whale. Endangered Species Research 14:39-48.

Kirk, Ruth. 1986. Tradition and change on the Northwest Coast: the Makah, Nuu-chah-nulth, southern Kwakiutl and Nuxalk. University of Washington Press, Seattle.

Lang, A. R., D. W. Weller, R. LeDuc, A. M. Burdin, V. L. Pease, D. Litovka, V. Burkanov, and R. L. Brownell, Jr. 2011. Genetic analysis of stock structure and movements of gray whales in the eastern and western North Pacific. SC/63/BRG10.

Moore, J. E. 2019. Declaration in re: ‘Proposed Waiver and Regulations Governing the Taking of Eastern North Pacific Gray Whales by the Makah Indian Tribe’. Administrative Law Judge, Hon. George J. Jordan. Docket No. 19-NMFS-0001. RINs: 0648-BI58; 0648-XG584.

Schubert, D. J. 2019. Rebuttal testimony in re: ‘Proposed Waiver and Regulations Governing the Taking of Eastern North Pacific Gray Whales by the Makah Indian Tribe’. Administrative Law Judge, Hon. George J. Jordan. Docket No. 19-NMFS-0001. RINs: 0648-BI58; 0648-XG584.

Scordino, J. J., M. Gosho, P. J. Gearin, A. Akmajian, J. Calambokidis, and N. Wright. 2017. Individual gray whale use of coastal waters off northwest Washington during the feeding season 1984-2011: Implications for management. Journal of Cetacean Research and Management 16:57-69.

Villegas-Amtmann, S. 2019. Declaration in re: ‘Proposed Waiver and Regulations Governing the Taking of Eastern North Pacific Gray Whales by the Makah Indian Tribe’. Administrative Law Judge, Hon. George J. Jordan. Docket No. 19-NMFS-0001.

Weller, D. W., S. Bettridge, R. L. Brownell, Jr., J. L. Laake, J. E. Moore, P. E. Rosel, B. L. Taylor, and P. R. Wade. 2013. Report of the National Marine Fisheries Service Gray Whale Stock Identification Workshop. NOAA-TM-NMFS-SWFSC-507. 

Yates, C. 2019a. Declaration in re: ‘Proposed Waiver and Regulations Governing the Taking of Eastern North Pacific Gray Whales by the Makah Indian Tribe’. Administrative Law Judge, Hon. George J. Jordan. Docket No. 19-NMFS-0001. RINs: 0648-BI58; 0648-XG584.

Yates, C. 2019b. Fifth declaration in re: ‘Proposed Waiver and Regulations Governing the Taking of Eastern North Pacific Gray Whales by the Makah Indian Tribe’. Administrative Law Judge, Hon. George J. Jordan. Docket No. 19-NMFS-0001. RINs: 0648-BI58; 0648-XG584.

The complex relationship between behavior and body condition

Clara Bird, Masters Student, OSU Department of Fisheries and Wildlife, Geospatial Ecology of Marine Megafauna Lab

Imagine that you are a wild foraging animal: In order to forage enough food to survive and be healthy you need to be healthy enough to move around to find and eat your food. Do you see the paradox? You need to be in good condition to forage, and you need to forage to be in good condition. This complex relationship between body condition and behavior is a central aspect of my thesis.

One of the great benefits of having drone data is that we can simultaneously collect data on the body condition of the whale and on its behavior. The GEMM lab has been measuring and monitoring the body condition of gray whales for several years (check out Leila’s blog on photogrammetry for a refresher on her research). However, there is not much research linking the body condition of whales to their behavior. Hence, I have expanded my background research beyond the marine world to looked for papers that tried to understand this connection between the two factors in non-cetaceans. The literature shows that there are examples of both, so let’s go through some case studies.

Ransom et al. (2010) studied the effect of a specific type of contraception on the behavior of a population of feral horses using a mixed model. Aside from looking at the effect of the treatment (a type of contraception), they also considered the effect of body condition. There was no difference in body condition between the treatment and control groups, however, they found that body condition was a strong predictor of feeding, resting, maintenance, and social behaviors. Females with better body condition spent less time foraging than females with poorer body condition. While it was not the main question of the study, these results provide a great example of taking into account the relationship between body condition and behavior when researching any disturbance effect.

While Ransom et al. (2010) did not find that body condition affected response to treatment, Beale and Monaghan (2004) found that body condition affected the response of seabirds to human disturbance. They altered the body condition of birds at different sites by providing extra food for several days leading up to a standardized disturbance. Then the authors recorded a set of response variables to a disturbance event, such as flush distance (the distance from the disturbance when the birds leave their location). Interestingly, they found that birds with better body condition responded earlier to the disturbance (i.e., when the disturbance was farther away) than birds with poorer body condition (Figure 1). The authors suggest that this was because individuals with better body condition could afford to respond sooner to a disturbance, while individuals with poorer body condition could not afford to stop foraging and move away, and therefore did not show a behavioral response. I emphasize behavioral response because it would have been interesting to monitor the vital rates of the birds during the experiment; maybe the birds’ heart rates increased even though they did not move away. This finding is important when evaluating disturbance effects and management approaches because it demonstrates the importance of considering body condition when evaluating impacts: animals that are in the worst condition, and therefore the individuals that are most vulnerable, may appear to be undisturbed when in reality they tolerate the disturbance because they cannot afford the energy or time to move away.

Figure 1.  Figure showing flush distance of birds that were fed (good body condition) and unfed (poor body condition).

These two studies are examples of body condition affecting behavior. However, a study on the effect of habitat deterioration on lizards showed that behavior can also affect body condition. To study this effect, Amo et al. (2007) compared the behavior and body condition of lizards in ski slopes to those in natural areas. They found that habitat deterioration led to an increased perceived risk of predation, which led to an increase in movement speed when crossing these deteriorated, “risky”, areas. In turn, this elevated movement cost led to a decrease in body condition (Figure 2). Hence, the lizard’s behavior affected their body condition.


Figure 2. Figure showing the difference in body condition of lizards in natural and deteriorated habitats.

Together, these case studies provide an interesting overview of the potential answers to the question: does body condition affect behavior or does behavior affect body condition? The answer is that the relationship can go both ways. Ransom et al. (2004) showed that regardless of the treatment, behavior of female horses differed between body conditions, indicating that regardless of a disturbance, body condition affects behavior. Beale and Monaghan (2004) demonstrated that seabird reactions to disturbance differed between body conditions, indicating that disturbance studies should take body condition into account. And, Amo et al. (2007) showed that disturbance affects behavior, which consequently affects body condition.

Looking at the results from these three studies, I can envision finding similar results in my gray whale research. I hypothesize that gray whale behavior varies by body condition in everyday circumstances and when the whale is disturbed. Yet, I also hypothesize that being disturbed will affect gray whale behavior and subsequently their body condition. Therefore, what I anticipate based on these studies is a circular relationship between behavior and body condition of gray whales: if an increase in perceived risk affects behavior and then body condition, maybe those affected individuals with poor body condition will respond differently to the disturbance. It is yet to be determined if a sequence like this could ever be detected, but I think that it is important to investigate.

Reading through these studies, I am ready and eager to start digging into these hypotheses with our data. I am especially excited that I will be able to perform this investigation on an individual level because we have identified the whales in each drone video. I am confident that this work will lead to some interesting and important results connecting behavior and health, thus opening avenues for further investigations to improve conservation studies.

References

Beale, Colin M, and Pat Monaghan. 2004. “Behavioural Responses to Human Disturbance: A Matter of Choice?” Animal Behaviour 68 (5): 1065–69. https://doi.org/10.1016/j.anbehav.2004.07.002.

Ransom, Jason I, Brian S Cade, and N. Thompson Hobbs. 2010. “Influences of Immunocontraception on Time Budgets, Social Behavior, and Body Condition in Feral Horses.” Applied Animal Behaviour Science 124 (1–2): 51–60. https://doi.org/10.1016/j.applanim.2010.01.015.

Amo, Luisa, Pilar López, and José Martín. 2007. “Habitat Deterioration Affects Body Condition of Lizards: A Behavioral Approach with Iberolacerta Cyreni Lizards Inhabiting Ski Resorts.” Biological Conservation 135 (1): 77–85. https://doi.org/10.1016/j.biocon.2006.09.020.

It all starts with the wind: The importance of upwelling

By Dawn Barlow, PhD student, OSU Department of Fisheries and Wildlife, Geospatial Ecology of Marine Megafauna Lab

The focus of my PhD research is on the ecology and distribution of blue whales in New Zealand. However, it has been a long time since I’ve seen a blue whale, and much of my time recently has been spent thinking about wind. What does wind matter to a blue whale? It actually matters a whole lot, because the wind drives an important biological process in many coastal oceans called upwelling. Wind blowing along shore, paired with the rotation of the earth, leads to a net movement of surface waters offshore (Fig. 1). As the surface water is pushed away, it is replaced by cold, nutrient-rich water from much deeper. When those nutrients become exposed to sunlight, they provide sustenance for the little planktonic lifeforms in the ocean, which in turn provide food for much larger predators including marine mammals such as blue whales. This “wind-to-whales” trophic pathway was coined by Croll et al. (2005), who demonstrated that off the West Coast of the United States, aggregations of whales could be expected downstream of upwelling centers, in concert with high productivity and abundant krill prey.

Figure 1. Graphic of the upwelling process, illustrating that when the wind blows along shore, surface waters are replaced by deeper water that is cold and nutrient rich. Source: NOAA
Figure 2. Map of New Zealand, with the South Taranaki Bight region (STB) denoted by the black box.

Much of what is understood today about upwelling comes from decades of research on the California Current ecosystem off the West Coast of the United States. Yet, the focus of my research is on an upwelling system on the other side of the world, in the South Taranaki Bight region (STB) of New Zealand (Fig. 2). In the case of the STB, westerly winds over Kahurangi Shoals lead to decreased sea level nearshore, forcing cold, nutrient rich waters to rise to the surface. The wind, along with the persistence of the Westland Current, then pushes a cold and productive plume of upwelled waters around Cape Farewell and into the STB (Fig. 3; Shirtcliffe et al. 1990).

Figure 3. Satellite image of the cold water plume in the South Taranaki Bight, indicative of upwelling. The origin of the upwelling at Kahurangi Shoals, Cape Farewell, and the typical path of the upwelling plume are denoted.

Through research conducted by the GEMM Lab over the years, we have demonstrated that blue whales utilize the STB region for foraging (Torres 2013, Barlow et al. 2018). Recent research on the oceanography of the STB region has further illuminated the mechanisms of this upwelling system, including the path and persistence of the upwelling plume in the STB across years and seasons (Chiswell et al. 2017, Stevens et al. 2019). However, the wind-to-whales pathway has not yet been described for this part of the world, and that is where the next section of my PhD research comes in. The whole system does not respond instantaneously to wind; the pathway from wind to whales takes time. But how much time is required for each step? How long after a strong wind event can we expect aggregations of feeding blue whales? These are some of the questions I am trying to tackle. For example, we hypothesize that some of the mechanisms and their respective lag times can be sketched out as follows:

Figure 4. The wind-to-whales trophic pathway, and hypothesized lags between steps.

All of these questions involve integrating oceanography, satellite imagery, wind data, and lag times, leading me to delve into many different analytical approaches including time series analysis and predictive modeling. If we are able to understand the lag times along this series of events leading to blue whale feeding opportunities, then we may be able to forecast blue whale occurrence in the STB based on the current wind and upwelling conditions. Forecasting with some amount of lead time could be a very powerful management tool, allowing for protection measures that are dynamic in space and time and therefore more effective in conserving this blue whale population and balancing human impacts.

Figure 5. A blue whale lunges on a patch of krill. The end of the wind-to-whales pathway. Drone piloted by Todd Chandler.

References:

Barlow DR, Torres LG, Hodge KB, Steel D, Baker CS, Chandler TE, Bott N, Constantine R, Double MC, Gill P, Glasgow D, Hamner RM, Lilley C, Ogle M, Olson PA, Peters C, Stockin KA, Tessaglia-hymes CT, Klinck H (2018) Documentation of a New Zealand blue whale population based on multiple lines of evidence. Endanger Species Res 36:27–40.

Chiswell SM, Zeldis JR, Hadfield MG, Pinkerton MH (2017) Wind-driven upwelling and surface chlorophyll blooms in Greater Cook Strait. New Zeal J Mar Freshw Res.

Croll DA, Marinovic B, Benson S, Chavez FP, Black N, Ternullo R, Tershy BR (2005) From wind to whales: Trophic links in a coastal upwelling system. Mar Ecol Prog Ser 289:117–130.

Shirtcliffe TGL, Moore MI, Cole AG, Viner AB, Baldwin R, Chapman B (1990) Dynamics of the Cape Farewell upwelling plume, New Zealand. New Zeal J Mar Freshw Res 24:555–568.

Stevens CL, O’Callaghan JM, Chiswell SM, Hadfield MG (2019) Physical oceanography of New Zealand/Aotearoa shelf seas–a review. New Zeal J Mar Freshw Res.

Torres LG (2013) Evidence for an unrecognised blue whale foraging ground in New Zealand. New Zeal J Mar Freshw Res 47:235–248.