Tag: whaling

A smaller sized gray whale: recent publication finds PCFG whales are smaller than ENP whales

Dr. KC Bierlich, Postdoctoral Scholar, OSU Department of Fisheries, Wildlife, & Conservation Sciences, Geospatial Ecology of Marine Megafauna (GEMM) Lab

A recent blog post by GEMM Lab’s PhD Candidate Clara Bird gave a recap of our 8th consecutive GRANITEfield season this year. In her blog, Clara highlighted that we saw 71 individual gray whales this season, 61 of which we have seen in previous years and identified as belonging to the Pacific Coast Feeding Group (PCFG). With an estimated population size of around 212 individuals, this means that we saw almost 1/3 of the PCFG population this season alone. Since the GEMM Lab first started collecting data on PCFG gray whales in 2016, we have collected drone imagery on over 120 individuals, which is over half the PCFG population. This dataset provides incredible opportunity to get to know these individuals and observe them from year to year as they grow and mature through different life history stages, such as producing a calf. A question our research team has been interested in is what makes a PCFG whale different from an Eastern North Pacific (ENP) gray whale, which has a population size around 16,000 individuals and feed predominantly in the Arctic during the summer months? For this blog, I will highlight findings from our recent publication in Biology Letters (Bierlich et al., 2023) comparing the morphology (body length, skull, and fluke size) between PCFG and ENP populations. 

Body size and shape reflect how an animal functions in their environment and can provide details on an individual’s current health, reproductive status, and energetic requirements. Understanding how animals grow is a key component for monitoring the health of populations and their vulnerability to climate change and other stressors in their environment.  As such, collecting accurate morphological measurements of individuals is essential to model growth and infer their health. Collecting such morphological measurements of whales is challenging, as you cannot ask a whale to hold still while you prepare the tape measure, but as discussed in a previous blog, drones provide a non-invasive method to collect body size measurements of whales. Photogrammetry is a non-invasive technique used to obtain morphological measurements of animals from photographs. The GEMM Lab uses drone-based photogrammetry to obtain morphological measurements of PCFG gray whales, such as their body length, skull length (as snout-to-blowhole), and fluke span (see Figure 1). 

Figure 1. Morphological measurements obtained via photogrammetry of a Pacific Coast Feeding Group (PCFG) gray whale. These measurements were used to compare to individuals from the Eastern North Pacific (ENP) population. 

As mentioned in this previous blog, we use photo-identification to identify unique individual gray whales based on markings on their body. This method is helpful for linking all the data we are collecting (morphology, hormones, behavior, new scarring and skin conditions, etc.) to each individual whale. An individual’s sightings history can also be used to estimate their age, either as a ‘minimum age’ based on the date of first sighting or a ‘known age’ if the individual was seen as a calf. By combining the length measurements from drone-based photogrammetry and age estimates from photo-identification history, we can construct length-at-age growth models to examine how PCFG gray whales grow. While no study has previously examined length-at-age growth models specifically for PCFG gray whales, another study constructed growth curves for ENP gray whales using body length and age estimates obtained from whaling, strandings, and aerial photogrammetry (Agbayani et al., 2020). For our study, we utilized these datasets and compared length-at-age growth, snout-to-blowhole length, and fluke span between PCFG and ENP whales. We used Bayesian statistics to account and incorporate the various levels of uncertainty associated with data collected (i.e., measurements from whaling vs. drone, ‘minimum age’ vs. ‘known age’). 

We found that while both populations grow at similar rates, PCFG gray whales reach smaller adult lengths than ENP. This difference was more extreme for females, where PCFG females were ~1 m (~3 ft) shorter than ENP females and PCFG males were ~0.5 m (1.5 ft) shorter than ENP males (Figure 2, Figure 3). We also found that ENP males and females have slightly larger skulls and flukes than PCFG male and females, respectively. Our results suggest PCFG whales are shaped differently than ENP whales (Figure 3)! These results are also interesting in light of our previous published study that found PCFG whales are skinnier than ENP whales (see this previous blog post). 

Figure 2. Growth curves (von Bertalanffy–Putter) for length-at-age comparing male and female ENP and PCFG gray whales (shading represents 95% highest posterior density intervals). Points represent mean length and median age. Vertical bars represent photogrammetric uncertainty. Dashed horizontal lines represent uncertainty in age estimates.

Figure 3. Schematic highlighting the differences in body size between Pacific Coast Feeding Group (PCFG) and Eastern North Pacific (ENP) gray whales. 

Our results raise some interesting questions regarding why PCFG are smaller: Is this difference in size and shape normal for this population and are they healthy? Or is this difference a sign that they are stressed, unhealthy and/or not getting enough to eat? Larger individuals are typically found at higher latitudes (this pattern is called Bergmann’s Rule), which could explain why ENP whales are larger since they feed in the Arctic. Yet many species, including fish, birds, reptiles, and mammals, have experienced reductions in body size due to changes in habitat and anthropogenic stressors (Gardner et al., 2011). The PCFG range is within closer proximity to major population centers compared to the ENP foraging grounds in the Arctic, which could plausibly cause increased stress levels, leading to decreased growth. 

The smaller morphology of PCFG may also be related to the different foraging tactics they employ on different prey and habitat types than ENP whales. Animal morphology is linked to behavior and habitat (see this blogpost). ENP whales feeding in the Arctic generally forage on benthic amphipods, while PCFG whales switch between benthic, epibenthic and planktonic prey, but mostly target epibenthic mysids. Within the PCFG range, gray whales often forage in rocky kelp beds close to shore in shallow water depths (approx. 10 m) that are on average four times shallower than whales feeding in the Arctic. The prey in the PCFG range is also found to be of equal or higher caloric value than prey in the Arctic range (see this blog), which is interesting since PCFG were found to be skinnier.

It is also unclear when the PCFG formed? ENP and PCFG whales are genetically similar, but photo-identification history reveals that calves born into the PCFG usually return to forage in this PCFG range, suggesting matrilineal site fidelity that contributes to the population structure. PCFG whales were first documented off our Oregon Coast in the 1970s (Figure 4). Though, from examining old whaling records, there may have been PCFG gray whales foraging off the coasts of Northern California to British Columbia since the 1920s.

Figure 4. First reports of summer-resident gray whales along the Oregon coast, likely part of the Pacific Coast Feeding Group. Capital Journal, August 9, 1976, pg. 2.

Altogether, our finding led us to two hypotheses: 1) the PCFG range provides an ecological opportunity for smaller whales to feed on a different prey type in a shallow environment, or 2) the PCFG range is an ecological trap, where individuals gain less energy due to energetically costly feeding behaviors in complex habitat while potentially targeting lower density prey, causing them to be skinnier and have decreased growth. Key questions remain for our research team regarding potential consequences of the smaller sized PCFG whales, such as does the smaller body size equate to reduced resilience to environmental and anthropogenic stressors? Does smaller size effect fecundity and population fitness? Stay tuned as we learn more about this unique and fascinating smaller sized gray whale. 

References

Agbayani, S., Fortune, S. M. E., & Trites, A. W. (2020). Growth and development of North Pacific gray whales (Eschrichtius robustus). Journal of Mammalogy, 101(3), 742–754. https://doi.org/10.1093/jmammal/gyaa028

Bierlich, K. C., Kane, A., Hildebrand, L., Bird, C. N., Fernandez Ajo, A., Stewart, J. D., Hewitt, J., Hildebrand, I., Sumich, J., & Torres, L. G. (2023). Downsized: gray whales using an alternative foraging ground have smaller morphology. Biology Letters, 19(8). https://doi.org/10.1098/rsbl.2023.0043

Gardner, J. L., Peters, A., Kearney, M. R., Joseph, L., & Heinsohn, R. (2011). Declining body size: A third universal response to warming? Trends in Ecology and Evolution, 26(6), 285–291. https://doi.org/10.1016/j.tree.2011.03.005

A Multidisciplinary Treasure Hunt: Learning about Indigenous Whaling in Oregon

By Rachel Kaplan, PhD student, OSU College of Earth, Ocean, and Atmospheric Sciences and Department of Fisheries and Wildlife, Geospatial Ecology of Marine Megafauna Lab

At this year’s virtual State of the Coast conference, I enjoyed tuning into a range of great talks, including one by Zach Penney from the Columbia River Inter-Tribal Fish Commission. In his presentation, “More Than a Tradition: Treaty rights and the Columbia River Inter-Tribal Fish Commission,” Penney described a tribal “covenant with resources,” and noted the success of this approach — “You don’t live in a place for 15,000 years by messing it up.”

Indigenous management of resources in the Pacific Northwest dates back thousands of years. From oak savannahs to fisheries to fires, local tribes managed diverse natural systems long before colonial settlement of the area that is now Oregon. We know comparatively little, however, about how Indigenous groups in Oregon interacted with whale populations before the changes brought by colonialism and commercial whaling.

Makah hunters in Washington bring a harvested whale into Neah Bay (Asahel Curtis/Washington State Historical Society).

I’m curious about how this missing knowledge could inform our understanding of the coastal Oregon ecosystems in which many GEMM Lab projects take place. My graduate research will be part of the effort to identify co-occurrence between whales and fishing in Oregon, with the goal of helping to reduce whale entanglement risk. Penney’s talk, ongoing conversations about decolonizing science, and my own concerns about becoming the scientist that I want to be, have all led me to ask a new set of questions: What did humans know in the past about whale distributions along the Oregon coast? What lost knowledge can be reclaimed from history?

As I started reading about historical Indigenous whale use in Oregon, I was struck by how little we know today, and how this learning process became a multidisciplinary treasure hunt. Clues as to how Indigenous groups interacted with whales along the Oregon coast lie in oral histories, myths, journals, and archaeological artifacts. 

Much of what I read hinged on the question: did Indigenous tribes in Oregon historically hunt whales? Many signs point to yes, but it’s a surprisingly tricky question to answer conclusively. Marine systems and animals, including seals and whales, remain an important part of cultures in the Pacific Northwest today – but historically, documentation of hunting whales in Oregon has been limited. Whale bones have been found in coastal middens, and written accounts describe opportunistic harvests of beached whales. However, people have long believed that only a few North American tribes outside of the Arctic regularly hunted whales. 

But in 2007, archaeologists Robert Losey and Dongya Yang found an artifact that started to shift this narrative. While studying a collection of tools housed at the Smithsonian Institution, they discovered the tip of a harpoon lodged in a whale flipper bone. This artifact came from the Partee site, which was inhabited around AD 300-1150 and is located near present-day Seaside, Oregon.

A gray whale ulna with cut marks found at the Partee site (Wellman, et al. 2017).

Through DNA testing, Losey and Yang determined that the harpoon was made of elk bone, and that the elk was not only harvested locally, but also used locally. This new piece of evidence suggested that whaling did in fact take place at the Partee site, likely by the Tillamook or Clatsop tribes that utilized this area.

Several years later, this discovery inspired Smithsonian Museum of Natural History archaeologist Torben Rick and University of Oregon PhD student Hannah Wellman to comb through the rest of the animal remains in the Smithsonian’s collection from northwest Oregon. Rick and Wellman scrutinized 187 whale bones for signs of hunting or processing, and found that about a quarter of the marks they inspected could have come from either hunting or the opportunistic harvest of stranded whales. They examined tools from the midden as well, and found that they were more suited to hunting animals, like seals and sea lions, or fishing. 

However, Wellman and Rick also used DNA testing to identify which whale species were represented in the midden – and the DNA analyses suggested a different story. Genetic results revealed that the majority of whale bones in the midden came from gray whales, a third from humpback whales, and a few from orca and minke. Modern gray whale stranding events are not uncommon, and so it follows logically that these bones could have simply come from people harvesting beached whales. However, humpback strandings are rare – suggesting that such a large proportion of humpback bones in the midden is likely evidence of people actively hunting humpback whales.

Percentage of whale species identified at the Partee site and percentage of species in the modern stranding record for the Oregon Coast (Wellman, et al. 2017).

These results shed new light on whale harvesting practices at the Partee Site, and, like so much research, they suggest a new set of questions. What does the fact that there were orca, minke, gray, and humpback whales off the Oregon coast 900 years ago tell us about the history of this ecosystem? Could artifacts that have not yet been found provide more conclusive evidence of hunting? What would it mean if these artifacts are found one day, or if they are never found?

As this fascinating research continues, I hope that new discoveries will continue to deepen our understanding of historic Indigenous whaling practices in Oregon – and that this information can find a place in contemporary conversations. Indigenous whaling rights are both a contemporary and contentious issue in the Pacific Northwest, and the way that humans learn about the past has much to do with how we shape the present. 

What we learn about the past can also change how we understand this ecosystem today, and provide new context as we try to understand the impacts of climate change on whale populations in Oregon. I’m interested in how learning more about historical Indigenous whaling practices could provide more information about whale population baselines, ideas for management strategies, and a new lens on the importance of whales in the Pacific Northwest. Even if we can’t fully reclaim lost knowledge from history, maybe we can still read enough clues to help us see both the past and present more fully.

Sources:

Braun, Ashley. “New Research Offers a Wider View on Indigenous North American Whaling.” Hakai Magazine, November 2016, www.hakaimagazine.com/news/new-research-offers-wider-view-indigenous-north-american-whaling/. 

Eligon, John. “A Native Tribe Wants to Resume Whaling. Whale Defenders Are Divided.” New York Times, November 2019. 

Hannah P. Wellman, Torben C. Rick, Antonia T. Rodrigues & Dongya Y. Yang (2017) Evaluating Ancient Whale Exploitation on the Northern Oregon Coast Through Ancient DNA and Zooarchaeological Analysis, The Journal of Island and Coastal Archaeology, 12:2, 255-275, DOI: 10.1080/15564894.2016.1172382

Losey, R., & Yang, D. (2007). Opportunistic Whale Hunting on the Southern Northwest Coast: Ancient DNA, Artifact, and Ethnographic Evidence. American Antiquity, 72(4), 657-676. doi:10.2307/25470439

Sanchez, Gabriel (2014). Conference paper: Cetacean Hunting at the Par-Tee site (35CLT20)?: Ethnographic, Artifact and Blood Residue Analysis Investigation.

Whales are amazing, and also provide amazing benefits to our oceans and human society

By Alejandro Fernandez Ajo, PhD student at the Department of Biology, Northern Arizona University, Visiting scientist in the GEMM Lab working on the gray whale physiology and ecology project  

Whales are among the most amazing and enigmatic animals in the world. Whales are not only fascinating, they are also biologically special. Due to their key ecological role and unique biological traits (i.e., their large body size, long lifespans, and sizable home ranges), whales are extremely important in helping sustain the entire marine ecosystem.

Working towards the conservation of marine megafauna, and large charismatic animals in general, is often seen as a mere benevolent effort that conservationist groups, individuals, and governments do on behalf of the individual species. However, mounting evidence demonstrates that restoring populations of marine megafauna, including large whales, can help buffer marine ecosystems from destabilizing stresses like human driven CO2 emissions and global change due to their ability to sequester carbon in their bodies (Pershing et al. 2010). Furthermore, whales can enhance primary production in the ocean through their high consumption and defecation rates, which ultimately provides nutrients to the ecosystem and improves fishery yields (Roman-McCarthy, 2010; Morissette et al. 2012).

Relationships between humans and whales have a long history, however, these relationships have changed. For centuries, whales were valued in terms of the number of oil barrels they could yield, and the quality of their baleen and meat. In the North Atlantic, whaling started as early as 1000 AD with “shore whaling” of North Atlantic right whales by Basque whalers. This whaling was initially limited to the mother and calve pairs that were easy to target due to their coastal habits and the fact that calves are more vulnerable and slower (Reeves-Smith, 2006). Once the calving populations of near-shore waters off Europe were depleted, offshore whaling began developing. Whalers of multiple nations (including USA, British, French, Norwegian, Portuguese, and Dutch, among others), targeted whales around the world, mainly impacting the gray whale populations, and all three right whale species along with the related bowhead whale. Later, throughout the phase of modern whaling using industrialized methods, the main target species consisted of the blue, fin, humpback, minke, sei and sperm whale (Schneider- Pearce, 2004).

By the early twentieth century, many of the world´s whale populations where reduced to a small fraction of their historical numbers, and although pre-whaling abundance of whale stocks is a subject of debate, recent studies estimate that at least the 66%, and perhaps as high as 90% for some whale species and populations (Branch-Williams 2006; Christensen, 2006), where taken during this period. This systematic and serial depletion of whale papulations reduced the biomass and abundance of great whales around the world, which has likely altered the structure and function of the oceans (Balance et al. 2006; Roman et al. 2014; Croll, et al. 2006).

After centuries of unregulated whale hunting, commercial whaling was banned in the mid-twentieth century. This ban was the result of multiple factors including reduced whale stocks below the point where commercial whaling would be profitable, and a fortunate shift in public perception of whales and the emergence of conservation initiatives (Schneider- Pearce, 2004). Since this moratorium on whaling, several whale populations have recovered around the world, and some populations that were listed as endangered have been delisted (i.e., the Eastern North Pacific gray whale) and some populations are estimated to have re-bounced to their pre-whaling abundance.

Although, the recovery of some populations has motivate some communities or nations to obtain or extend their whaling quotas (see Blog Post by Lisa Hildebrand), it is important to acknowledge that the management of whale populations is arguably one of the most complicated tasks, and is distinguished from management of normal fisheries due to various biological aspects. Whales are long living mammals with slow reproduction rates, and on average a whale can only produce a calf every two or three years. Hence, the gross addition to the stock rarely would exceed 25% of the number of adults (Schneider- Pearce, 2004), which is a much lower recovery rate that any fish stock. Also, whales usually reach their age of sexual maturity at 6-10 years old, and for many species there are several uncertainties about their biology and natural history that make estimations of population abundance and growth rate even harder to estimate.

Fig 1: Human relationship with whales has changed through history. Once valued for their meat and oil, now they are a natural attraction that amaze and attract crowds to whale watching destinations all over the globe. Photo: Stephen Johnson, Península Valdés-Argentina.

Moreover, while today´s whales are generally not killed directly by hunting, they are exposed to a variety of other increasing human stressors (e.g., entanglement in fishing gear, vessel strikes, shipping noise, and climate change). Thus, scientists must develop novel tools to overcome the challenges of studying whales and distinguish the relative importance of the different impacts to help guide conservation actions that improve the recovery and restoration of whale stocks (Hunt et al. in press). With the restoration of great whale populations, we can expect positive changes in the structure and function of the world’s oceans (Chami et al. 2019; Roman et al. 2010).

So, why it is worth keeping whales healthy?

Whales facilitate the transfer of nutrients by (1) releasing nutrient-rich fecal plumes near the surface after they have feed at depth and (2) by moving nutrients from highly productive, polar and subpolar latitude feeding areas to the low latitude calving areas (Roman et al. 2010). In this way, whales help increase the productivity of phytoplankton that in turn support zooplankton production, and thus have a bottom up effect on the productivity of many species including fish, birds, and marine mammals, including whales. These fertilization events can also facilitate mitigation of the negative impacts of climate change. The amount of iron contained in the whales’ feces can be 10 million times greater than the level of iron in the marine environment, triggering important phytoplankton blooms, which in turn sequester thousands of tons of carbon from, and release oxygen to, the atmosphere annually (Roman et al. 2016; Smith et al. 2013; Willis, 2007). Furthermore, when whales die, their massive bodies fall to the seafloor, making them the largest and most nutritious source of food waste, which is capable of sustaining a succession of macro-fauna assemblages for several decades, including some invertebrate species that are endemic to whale carcasses (Smith et al. 2015).

Figure 2. The figure shows a conceptual model of the “whale pump”. From Roman-McCarthy, 2010.

Despite the several environmental services that whales provide, and the positive impact on local economies that depend on whale watching tourism, which has been valued in millions of dollars per year (Hoyt E., 2001), the return of whales and other marine mammals has often been implicated in declines in fish populations, resulting in conflicts with human fisheries (Lavigne, D.M. 2003). Yet there is insufficient direct evidence for such competition (Morissette et al. 2010). Indeed, there is evidence of the contrary: In ecosystem models where whale abundances are reduced, fish stocks show significant decreases, and in some cases the presence of whales in these models result in improved fishery yields. Consistent with these findings, several models have shown that alterations in marine ecosystems resulting from the removal of whales and other marine mammals do not lead to increases in human fishery yields (Morissette et al. 2010; 2012). Although the environmental services and benefits provided by great whales, which potentially includes the enhancement of fisheries yields, and enhancement on ocean oxygen production and capturing carbon, are evident and make a strong argument for improved whale conservation, it is overwhelming how little we know about many aspects of their lives, their biology, and particularly their physiology.

Figure 3: Whales are the most fascinating animals in the world, but they are not only amazing animals. They are also extremely important in sustaining the entire marine ecosystem. Photo: Alejandro Fernández Ajó -Instituto de Conservación de ballenas.

This lack of knowledge is because whales are really hard to study. For many years research was limited to the observation of the brief surfacing of the whales, yet most of their lives occurs beneath the surface and were completely unknown. Fortunately, new technologies and the creativity of whale researchers are helping us to better understand many aspects of their lives that were cryptic to us even a decade ago. I am committed to filling some of these knowledge gaps. My research examines how different environmental and anthropogenic impacts affect whale health, and particularly how these impacts may relate to cases of large whale mortalities and declines in whale populations. I am applying novel methods in conservation physiology for measuring hormone levels that promise to improve our understanding of the relationship between different (extrinsic and intrinsic) stressors and the physiological response of whales. Ultimately, this research will help address important conservation questions, such as the causes of unusual whale mortality events and declines in whale populations.

References:

Ballance LT, Pitman RL, Hewitt R, et al. 2006. The removal of large whales from the Southern Ocean: evidence for long-term ecosystem effects. In: Estes JA, DeMaster DP, Doak DF, et al. (Eds). Whales, whaling and ocean ecosystems. Berkeley, CA: University of California Press.

Branch TA and Williams TM. 2006. Legacy of industrial whaling. In: Estes JA, DeMaster DP, Doak DF, et al. (Eds). Whales, whaling and ocean ecosystems. Berkeley, CA: University of California Press.

Chami, R. Cosimano, T. Fullenkamp, C. & Oztosun, S. (2019). Nature’s solution to climate change. Finance & Development, 56(4).

Christensen LB. 2006. Marine mammal populations: reconstructing historical abundances at the global scale. Vancouver, Canada: University of British Columbia.

Croll DA, Kudela R, Tershy BR (2006) Ecosystem impact of the decline of large whales in the North Pacific. In: Estes JA, DeMaster DP, Doak DF, Williams TM, BrownellJr RL, editors. Whales, Whaling, and Ocean Ecosystems. Berkeley: University of California Press. pp. 202–214.

Hoyt, E. 2001. Whale Watching 2001: Worldwide Tourism Numbers, Expenditures and Expanding Socioeconomic Benefits

Hunt, K.E., Fernández Ajó, A. Lowe, C. Burgess, E.A. Buck, C.L. In press. A tale of two whales: putting physiological tools to work for North Atlantic and southern right whales. In: “Conservation Physiology: Integrating Physiology Into Animal Conservation And Management”, ch. 12. Eds. Madliger CL, Franklin CE, Love OP, Cooke SJ. Oxford University press: Oxford, UK.

Lavigne, D.M. 2003. Marine mammals and fisheries: the role of science in the culling debate. In: Gales N, Hindell M, and Kirkwood R (Eds). Marine mammals: fisheries, tourism, and management issues. Melbourne, Australia: CSIRO.

Morissette L, Christensen V, and Pauly D. 2012. Marine mammal impacts in exploited ecosystems: would large scale culling benefit fisheries? PLoS ONE 7: e43966.

Morissette L, Kaschner K, and Gerber LR. 2010. “Whales eat fish”? Demystifying the myth in the Caribbean marine ecosystem. Fish Fish 11: 388–404.

Pershing AJ, Christensen LB, Record NR, Sherwood GD, Stetson PB (2010) The impact of whaling on the ocean carbon cycle: Why bigger was better. PLoS ONE 5(8): e12444.

Reeves, R. and Smith, T. (2006). A taxonomy of world whaling. In DeMaster, D. P., Doak, D. F., Williams, T. M., and Brownell Jr., R. L., eds. Whales, Whaling, and Ocean Ecosystems. University of California Press, Berkeley, CA.

Roman, J. Altman I, Dunphy-Daly MM, et al. 2013. The Marine Mammal Protection Act at 40: status, recovery, and future of US marine mammals. Ann NY Acad Sci; doi:10.1111/nyas.12040.

Roman, J. and McCarthy, J.J. 2010. The whale pump: marine mammals enhance primary productivity in a coastal basin. PLoS ONE. 5(10): e13255.

Roman, J. Estes, J.A. Morissette, L. Smith, C. Costa, D. McCarthy, J. Nation, J.B. Nicol, S. Pershing, A.and Smetacek, V. 2014. Whales as marine ecosystem engineers. Frontiers in Ecology and the Environment. 12(7). 377-385.

Roman, J. Nevins, J. Altabet, M. Koopman, H. and McCarthy, J. 2016. Endangered right whales enhance primary productivity in the Bay of Fundy. PLoS ONE. 11(6): e0156553.

Schneider, V. Pearce, D. What saved the whales? An economic analysis of 20th century whaling. Biodiversity and Conservation 13, 543–562 (2004). https://doi org.libproxy.nau.edu/10.1023/B:BIOC.0000009489.08502.1

Smith LV, McMinn A, Martin A, et al. 2013. Preliminary investigation into the stimulation of phyto- plankton photophysiology and growth by whale faeces. J Exp Mar Biol Ecol 446: 1–9.

Smith, C.R. Glover, A.G. Treude, T. Higgs, N.D. and Amon, D.J. 2015. Whale-fall ecosystems: Recent insights into ecology, paleoecology, and evolution. Annu. Rev. Marine. Sci. 7:571-596.

Willis, J. 2007. Could whales have maintained a high abundance of krill? Evol Ecol Res 9: 651–662.