Extreme Cetaceans, Part 1

It was while going through my read all the books on the whales of the world phase of the early 1990s, I remember, that I first read of the dolphins – members of the highly streamlined, long-beaked, oceanic dolphin group no less – that have such weird features as deep keels, humps on the back and tailstock, and non-streamlined, forward-canted dorsal fins. Yes, we all know that whales are streamlined, torpedo-shaped animals with sensibly shaped appendages, but they’re not all like this. Quite a few species are weird, possessing anatomical specialisations and peculiarities that are counter-intuitive and little discussed, and most likely related to an unusual ecology, physiological regime, feeding strategy or social or sexual life.

A nice, normal looking group of Spinner dolphins. The obvious dark cape and paler side regions make these look like Hawaiian spinners but they were apparently photographed in the Red Sea. Image: Alexander Vasenin, CC BY-SA 3.0, wikipedia (original h…

Caption: a nice, normal looking group of Spinner dolphins. The obvious dark cape and paler side regions make these look like Hawaiian spinners but they were apparently photographed in the Red Sea. Image: Alexander Vasenin, CC BY-SA 3.0, wikipedia (original here).

In this short series of articles – yeah, this is Part 1 – I want to talk about just a few such animals, and I hope you’ll be as surprised by their anatomy and specialisations as I was when I first learnt about them.

Extreme spinners. Everybody knows that dolphins are streamlined, and the oceanic long-beaked dolphins (those conventionally united in the genus Stenella) are streamlined the most. The Spinner S. longirostris – a species that occurs throughout the tropical and subtropical marine waters of the world – is one such animal, its very long beak, torpedo-shaped body and tailstock and well-proportioned fins all appearing like textbook adaptations for swift movement in the pelagic environment. Yet for all this, some spinner dolphins – some specific individuals belonging to some specific populations – are very odd indeed.

One of the very best depictions of an ‘extreme’ male Eastern spinner is this one, from Shirahai & Jarrett’s 2006 Whales, Dolphins and Seals. Image: (c) Brett Jarrett.

Caption: one of the very best depictions of an ‘extreme’ male Eastern spinner is this one, from Shirahai & Jarrett’s 2006 Whales, Dolphins and Seals. Image: (c) Brett Jarrett.

These animals have arching dorsal humps and massive, bulbous ventral convexities on the tailstock which give them a peculiarly asymmetrical, lumpy appearance, the dorsal fin is not recurved and falcate, but has a straight or even concave anterior margin such that it might even lean forwards, and the tail flukes turn upwards at their outer edges. The exaggerated lump on the lower surface of the tailstock has a name: it’s the post-anal hump. This structure isn’t unique to the Spinner but is also present in other delphinids, like the Delphinus species. It appears to be sexually dimorphic and is especially prominent in mature males (Ngqulana et al. 2017). Perrin & Mesnick (2003) argued that these features - which are variable in spinner populations and most strongly developed in the so-called Eastern and Whitebelly spinners - are linked to testis size and to a polygynous mating system where males need to be highly distinct from their many female consorts, and built to display against, and fight with, other males. In other words, the most ‘extreme’ spinners are the most polygynous.

Adult males differ in appearance across spinner populations, and it seems that the most ‘extreme’ males are those from the most polygynous populations. This diagram (from Perrin & Mesnick 2003) shows - from top to bottom - male Hawaiian, Eastern…

Caption: adult males differ in appearance across spinner populations, and it seems that the most ‘extreme’ males are those from the most polygynous populations. This diagram (from Perrin & Mesnick 2003) shows - from top to bottom - male Hawaiian, Eastern and ‘whitebelly’ spinners. Image: Perrin & Mesnick (2003).

How and why might this remarkable feature have originated? Spinners and other cetaceans adopt a sinuous, vaguely S-shaped profile when displaying to one another (this has now been seen in diverse cetaceans, mysticetes as well as odontocetes; Helweg et al. 1992, Horback et al. 2010), and one suggestion is that the post-anal hump and a matching convexity on the dorsal surface of the tailstock might serve to accentuate the curves of the S and thereby function in exaggerating this signal. One idea about the S-shaped pose is that it functions in shark mimicry (Norris et al. 1985; some sharks also adopt an S-shaped profile and use it to signal aggressive intentions), but the fact that it’s as widespread in cetaceans as it is – and similar poses are seen in other aquatic vertebrates – indicates that any similarities with non-cetaceans are convergent.

S-shaped postures, depicted (sometimes schematically) in cetaceans of very different sizes and proportions, from Horback et al. (2010). (A) Spinner dolphin, (B) Beluga, (C) Humpback whale. Evolve dorsal and ventral convexities on the body and tailst…

Caption: S-shaped postures, depicted (sometimes schematically) in cetaceans of very different sizes and proportions, from Horback et al. (2010). (A) Spinner dolphin, (B) Beluga, (C) Humpback whale. Evolve dorsal and ventral convexities on the body and tailstock, and you can exaggerate the intensity of this signal. Image: Horback et al. (2010).

Anyway… the features discussed here appear intuitively odd because they’re just about the opposite of what you’d predict to be present in a fast-swimming, pelagic predator which has otherwise evolved to be ultra-streamlined. But there you are.

Humpback dolphins are not especially well known, and even less well known is that they’re kept in captivity in a few places and have been trained to do tricks. This individual was photographed in captivity in Singapore. Image: Tolomea, CC BY 2.0, wi…

Caption: humpback dolphins are not especially well known, and even less well known is that they’re kept in captivity in a few places and have been trained to do tricks. This individual was photographed in captivity in Singapore. Image: Tolomea, CC BY 2.0, wikipedia (original here).

The humpback dolphins. Everyone’s heard of the Humpback whale Megaptera novaeangliae, but less well known is that there are dolphins with humps too, perhaps four species of them if you follow some studies of molecular variation within the group (the Indo-Pacific humpback dolphin Sousa chinensis, Australian humpback dolphin S. sahulensis, Atlantic humpback dolphin S. teuszii and Indian Ocean humpback dolphin S. plumbea). Superficially, Sousa dolphins look something like bottlenose dolphins (Tursiops), and like them they’re coastal animals that prey on diverse fishes and cephalopods. Unlike the Tursiops dolphins, the Sousa species have a long raised section – sitting dorsal to the neural spines and musculature of the back – that extends along the middle part of the dorsal surface. The dorsal fin sits on top of this hump.

Comparatively few people know that there are dolphins with humps, but check it out. These are Tom Ritchie’s illustrations of Sousa dolphins, representing adult males identified by Watson (1981) as S. chinensis (above) and S. teuszii (below). Images:…

Caption: comparatively few people know that there are dolphins with humps, but check it out. These are Tom Ritchie’s illustrations of Sousa dolphins, representing adult males identified by Watson (1981) as S. chinensis (above) and S. teuszii (below). Images: Watson 1981.

The function of this hump – if it has one – is not well studied and authors have mostly avoided mentioning the possibility that it might have one. Does it function as a visual or acoustic signal of maturity? Does it have some role in buoyancy, hydrodynamics or streamlining? Is it a fat store? The dorsal fins of at least some cetaceans appear to function as so-called thermal windows: as heat-dumping structures, the large and extensive blood vessels of which carry cooled blood to the body interior (Meagher et al. 2002). In males, this cool blood helps lower the temperature of the deeply internal testes (Pabst et al. 1995), which might otherwise be prone to overheating. The humps of humpbacked dolphins, like the dorsal fins, appear to be richly innervated with blood vessels which again transport cooled blood from the animal’s exterior surface to deep within its body (Plön et al. 2018).

(A) Vasculature in the dorsal fin and hump of a humpback dolphin compared with (B) dorsal fin vasculature in a Tursiops dolphin. The blood vessels in Tursiops are proportionally larger, but there’s a great number of them in the humpback dolphin, tha…

Caption: (A) vasculature in the dorsal fin and hump of a humpback dolphin compared with (B) dorsal fin vasculature in a Tursiops dolphin. The blood vessels in Tursiops are proportionally larger, but there’s a great number of them in the humpback dolphin, thanks to the hump. Image: Plön et al. 2018.

Could the hump therefore be a thermoregulatory specialisation for this mostly tropical group? Further research is needed, but this could be consistent with the fact that the hump is proportionally largest in adult males, and perhaps proportionally largest in those populations that inhabit the most tropical parts of Sousa’s entire range. A hydrodynamic role for the hump remains plausible but has yet to be investigated (Plön et al. 2018).

And that’s where we’ll end things for now. More in this series soon. I’ll publish a lot more on whales here in the future, but here’s some of the stuff that exists in the archives (as always, much of the material at TetZoo versions 2 and 3 has been ruined by the removal of images)…

If you enjoyed this article and want to see me do more, more often, please consider supporting me at patreon. The more funding I receive, the more time I’m able to devote to producing material for TetZoo and the more productive I can be on those long-overdue book projects. Thanks!

Refs - -

Helweg, D. A., Bauer, G. B. & Herman, L. M. 1992. Observations of an S-shaped posture in humpback whales (Megaptera novaeangliae). Aquatic Mammals 18.3, 74-78.

Horback, K. M., Friedman, W. R. & Johnson, C. M. 2010. The occurrence and context of S-posture display by captive belugas (Delphinapterus leucas). International Journal of Comparative Psychology 23, 689-700.

Meagher, E. M., McLellan, W. A., Westgate, A. J., Wells, R. S., Frierson, D. Jr. & Pabst, D. A.. 2002. The relationship between heat flow and vasculature in the dorsal fin of wild bottlenose dolphins Tursiops truncatus. Journal of Experimental Biology 205, 3475-3486.

Pabst, D. A., Rommel, S. A., McLellan, W. A., Williams, T. M. & Rowles, T. K. 1995. Thermoregulation of the intra-abdominal testes of the bottlenose dolphin (Tursiops truncatus) during exercise. Journal of Experimental Biology 198, 221-226.

Norris, K. S., Wursig, B., Wells, R. S., Wursig, M., Brownlee, S. M., Johnson, C. & Solow, J. 1985. Behavior of the Hawaiian spinner dolphin, Stenella longirostris. National Marine Fisheries Service Administrative Report LJ-85-06C.

Ngqulana, S. G., Hofmeyr, G. J. G. & Plön, S. 2017. Sexual dimorphism in long-beaked common dolphins (Delphinus capensis) from KwaZulu-Natal, South Africa. Journal of Mammalogy 98, 1389-1399.

Perrin, W. F. & Mesnick, S. L. 2003. Sexual ecology of the Spinner dolphin, Stenella longirostris: geographic variation in mating system. Marine Mammal Science 19, 462-483.

Plön, S., Frainer, G., Wedderburn-Maxwell, A., Cliff, G. & Huggenberger, S. 2018. Dorsal fin and hump vascular anatomy in the Indo-Pacific humpback dolphin (Sousa plumbea) and the Indo-Pacific bottlenose dolphin (Tursiops aduncus). Marine Mammal Science 35, 684-695.

Shirihai, H. & Jarrett, B. 2006. Whales, Dolphin and Seals: a Field Guide to the Marine Mammals of the World. A & C Black, London.

Watson, L. 1981. Whales of the World. Hutchinson, London.

Whale Watching in the Bay of Biscay

Back in July 2019, myself and a bunch of friends stepped aboard the Pont-Aven for several days of sea-watching in the Bay of Biscay. We were to travel from Plymouth (UK) to Santander (Spain), the event being organised by ORCA, a charity that monitors whales and uses the data for conservation purposes (they’re here on Twitter). ORCA uses cruise liners, ferries and other vehicles as whale-watching platforms. Nigel Marven was a special guest on our trip and it was great to catch up with him.

Our vessel of choice - the Pont-Aven - at port in Santander, Spain. I cannot tell you how much trouble I went to to get to this ship before departure time. I very nearly didn’t make it. Image: Darren Naish.

Caption: our vessel of choice - the Pont-Aven - at port in Santander, Spain. I cannot tell you how much trouble I went to to get to this ship before departure time. I very nearly didn’t make it. Image: Darren Naish.

The man, the legend; Nigel Marven.

Caption: the man, the legend; Nigel Marven.

The purpose, of course, was to see whales. The weather was outstandingly good (meaning that I got burnt), but so was the whale watching: I’m pleased to say that we saw literally hundreds of animals of seven or eight species, as you can see from the photos below. My own photos are not great since my camera isn’t exactly the best for fast-moving, far-away animals like whales, so those you see here were mostly taken by my trusty pal Alex Srdic (who’s here on Instagram and here on Twitter). Thanks, Alex.

Several cetaceans have extremely complex markings allowing them to be identified to species and even population. Individuals can be recognised on the basis of their markings too. Image: Alex Srdic.

Caption: several cetaceans have extremely complex markings allowing them to be identified to species and even population. Individuals can be recognised on the basis of their markings too. Image: Alex Srdic.

The Bay of Biscay is a world-famous whale-watching hotspot, famous in particular for Cuvier’s beaked whales Ziphius cavirostris and Sperm whale Physeter macrocephalus. Dolphins of several species are a frequent sight too, as are rorquals of a few species, Harbour porpoise Phocoena phocoena and pilot whales. A very lucky whale-watcher might get to see Blue whale Balaenoptera musculus, Killer whale Orcinus orca or True’s beaked whale Mesoplodon mirus. In fact, something like 30 species have been recorded in the region. This is phenomenal and mean that it’s theoretically possible for several species of some of the most elusive whale groups – like beaked whales and globicephaline dolphins – to be seen within days or weeks of each other.

In good weather, the blow of a big whale (like a Fin whale - as here - or a Sperm whale) is visible from great distance, and in the case of these two species can be diagnostic. Image: Alex Srdic.

Caption: in good weather, the blow of a big whale (like a Fin whale - as here - or a Sperm whale) is visible from great distance, and in the case of these two species can be diagnostic. Image: Alex Srdic.

A dynamic leap by a Striped dolphin. Dolphins of some species appear to be attracted to ships and even to deliberately show off when they get close to them. Image: Alex Srdic.

Caption: a dynamic leap by a Striped dolphin. Dolphins of some species appear to be attracted to ships and even to deliberately show off when they get close to them. Image: Alex Srdic.

Why is the Bay of Biscay so good for whales? It’s mostly because the topography is complex, combining large, shallow shelf regions, steep sections of shelf edge – sometimes with impressive slopes and deep, enormous rocky canyons twice as big as the Grand Canyon – and a deep abyssal plain section (Carwardine 2016). Depth varies from 1.7 to over 4.7 km. This variation – combined with the overall productivity of the region and its position relative to the Atlantic and English Channel – means that there’s the chance to see continental shelf species (like porpoises), those that use deep canyons and other shelf-edge habitats (like beaked whales) and true oceanic deep-divers that forage in the deepest waters (like sperm whales).

Back and dorsal fin of a Fin whale, remnants of the blow still hanging in the air. Image: Alex Srdic.

Caption: back and dorsal fin of a Fin whale, remnants of the blow still hanging in the air. Image: Alex Srdic.

As it happens, we were extraordinarily lucky. Fin whales B. physalus are regular animals of the area, and we had amazing, relatively close views of them (by ‘close’, I mean perhaps 30 m from the ship, not alongside the vessel). Fin whales – the second largest extant animal after the Blue – have a blow that’s visible on the horizon and is about 8 m tall. The blow hangs in the air for a surprising time. One of the most remarkable things about the Fin whale is its asymmetrical pigmentation: the right side of the face is marked with a large pale area, as is the right side’s baleen. There are some old TetZoo articles on what this might mean and how it might function – see the links below.

Excellent view of the splashguard - the conical structure surrounding and ahead of the blowholes - and paired blowholes of a surfacing Fin whale. Despite its name, the dorsal fin of the Fin whale is smaller and blunter than that of some other rorqua…

Caption: excellent view of the splashguard - the conical structure surrounding and ahead of the blowholes - and paired blowholes of a surfacing Fin whale. Despite its name, the dorsal fin of the Fin whale is smaller and blunter than that of some other rorquals. Image: Alex Srdic.

Two coastal species were seen early on in our trip: Harbour porpoise and Common bottlenose dolphins Tursiops truncatus, though I don’t have good photos of either. The majority of dolphins seen on our trip (as is typical for Biscay whale watching) were Short-beaked common dolphin Delphinus delphis, which were sometimes seen in groups of more than ten. Their distinctive hourglass markings are always visible when they leap – which they often do, sometimes while immediately adjacent to a ship – and we also got to see calves on one or two occasions.

Here’s the whole-body view of the common dolphin shown in detail above. This individual only has one stripe extending from the beak to the flipper, with a large pale area separating the eye and flipper. Different configurations are present in differ…

Caption: here’s the whole-body view of the common dolphin shown in detail above. This individual only has one stripe extending from the beak to the flipper, with a large pale area separating the eye and flipper. Different configurations are present in different populations. Image: Alex Srdic.

As the light begins to fade during the later part of the day, a group of Short-beaked common dolphin carve through a surging wave. Note the calf close to the adult at upper right. Image: Alex Srdic.

Caption: as the light begins to fade during the later part of the day, a group of Short-beaked common dolphin carve through a surging wave. Note the calf close to the adult at upper right. Image: Alex Srdic.

We also had excellent views of Striped dolphin Stenella coeruleoalba. They behaved in characteristic acrobatic fashion, leaping high out of the water, making impressive splashes and jumping in the ship’s wake. They typically make a lot more disturbance at the water’s surface than do common dolphins, creating great bursts of spray and rooster-tail patterns when they leap and surge. Striped dolphins are near-globally distributed. They’ve been the source of discussion lately since it’s recently been shown that the Clymene dolphin S. clymene is a naturally occurring hybrid between this species and the Spinner S. longirostris (Amaral et al. 2014).

We had many excellent views of high-leaping Striped dolphin. Note how much spray and splashing is associated with the leaping of this species. Image: Alex Srdic.

Caption: we had many excellent views of high-leaping Striped dolphin. Note how much spray and splashing is associated with the leaping of this species. Image: Alex Srdic.

Finally as goes dolphins, we also saw pilot whales, identified on the basis of their black colouration and strongly backswept dorsal fins. These were most likely Long-finned pilots Globicephala melas (it’s more typical of temperate and cold waters than the Short-finned G. macrorhynchus) but we didn’t see any of the key features that allow the two species to be distinguished, and none of our photos are good enough to warrant sharing. A mysterious whale was seen among the pilot whales. It seemed to be very dark and with a short, blunt-tipped, parallel-sided but only weakly curved dorsal fin; I don’t think that its head was seen but I had the impression that it was a shallower-bodied animal than the pilot whales. Several different views were offered on its identity with the most likely (on the basis of dorsal fin shape) being that it was perhaps a False killer whale Pseudorca crassidens. That’s not tremendously likely but not impossible.

The whale most famously associated with the Bay of Biscay is Cuvier’s beaked whale, seen so frequently in the area that it’s regarded as the premier location for sightings of this species, worldwide. I don’t know if you’re guaranteed a sighting of a Cuvier’s while there, but – whatever – we were lucky, since we saw nearly 20 of them, ranging from smooth, clean-bodied youngsters to heavily scarred males.

Cuvier’s beaked whale, seen relatively close to the ship. Image: Alex Srdic.

Caption: Cuvier’s beaked whale, seen relatively close to the ship. Image: Alex Srdic.

Heavily scarred Cuvier’s beaked whale, seen at distance and only briefly. We didn’t see any other individuals with scarring as impressive as this. Image: Alex Srdic.

Caption: heavily scarred Cuvier’s beaked whale, seen at distance and only briefly. We didn’t see any other individuals with scarring as impressive as this. Image: Alex Srdic.

Some individuals have markedly pale heads sharply demarcated from the rest of the body, others do not. On occasion, one or two individuals were close enough to the ship that I was able to get a half-decent shot with my mobile phone. Each sighting was a huge thrill. While we were oh so lucky as goes Cuvier’s, we didn’t see sperm whale, alas. We also saw Northern minke B. acutorostrata on perhaps two occasions, though again I don’t have any good photos.

Another plus: amazing sunsets, and sunrises too. Image: Darren Naish.

Caption: another plus… amazing sunsets, and sunrises too. Image: Darren Naish.

Finally, we didn’t just see whales. The same route is also great for seabirds, and we also saw such fishes as tunas and sunfishes. As much as I’d like to start talking about the birds, I’m out of time. Anyway – the trip was excellent: rewarding, fun, and educational. I’ll definitely be doing it again. You should consider supporting ORCA and their work as well.

If you enjoyed this article and want to see me do more, more often, please consider supporting me at patreon. The more funding I receive, the more time I’m able to devote to producing material for TetZoo and the more productive I can be on those long-overdue book projects. Thanks!

Cetaceans have been covered at length on TetZoo before - mostly at ver 2 and ver 3 - but these articles are now all but useless since all of their images have been removed (and/or they’re paywalled, thanks SciAm). Over time, I aim to build up a large number of cetacean-themed articles here at ver 4.

Refs - -

Amaral, A. R., Lovewell, G., Coelho, M. M., Amato, G. & Rosenbaum, H. C. 2014. Hybrid speciation in a marine mammal: the Clymene dolphin (Stenella clymene). PLoS ONE 9 (1): e83645.

Carwardine, M. 2016. Mark Carwardine’s Guide to Whale Watching in Britain and Europe. Bloomsbury, London.

Remembering Lyall Watson’s Whales of the World

I’ve written before about some of the books that had an undue influence on me during my formative years. Such books tend to be well illustrated, they mostly contain attractive, colourful, detailed pieces of art, and they usually showcase weird and surprising proposals and arguments that later proved erroneous, questionable or wrong. The fact that I’ve always considered such books especially interesting and/or influential surely says a lot about me and how my brain works, but whatever.

The somewhat worn cover of my copy of Watson’s Whales of the World (the 1988 softback edition). Image: Darren Naish.

Caption: the somewhat worn cover of my copy of Watson’s Whales of the World (the 1988 softback edition). Image: Darren Naish.

Today I’d like to discuss another of these fondly remembered books, and if you know it as well as I do you may well understand where I’m coming from. If you don’t know the book at all, (1) what have you been doing with your life?, and (2) obtain the book for yourself, it’s worth it. I’m here to discuss the weird, wonderful Whales of the World (also published as Sea Guide to Whales of the World) by the late Lyall Watson, illustrated by Tom Ritchie, and subtitled ‘A Complete Guide to the World’s Living Whales, Dolphins and Porpoises’ (Watson 1981).

Watson (1981) is a robust, attractively designed volume of 302 pages that goes through all the cetacean species thought valid by the author at the time of writing. It saw at least three reprintings, the first edition being hardback with a dustjacket, the 1985 and 1988 editions being softbacks. The book is arranged taxonomically and groups the cetaceans together by family, each family section including an introduction that has a key and a guide to the family’s respective anatomical traits. The family-level taxonomy Watson used is a little idiosyncratic, on which more later. Each species gets its own two pages. These include a distribution map, colour illustration (sometimes showing variants and juveniles where appropriate), an image of the skull where possible, and text sections on Classification (read: taxonomic history and discovery), Local Names, Description, Stranding, Natural History, Status, Distribution, and Sources (there’s a good bibliography).

Watson (1981) includes both ‘wet keys’ (providing information on the life appearance of cetaceans, and intended to be used in the field) and ‘dry keys’ (providing information on skeletal material meant to be used to identify stranded animals or carc…

Caption: Watson (1981) includes both ‘wet keys’ (providing information on the life appearance of cetaceans, and intended to be used in the field) and ‘dry keys’ (providing information on skeletal material meant to be used to identify stranded animals or carcasses). Image: Watson 1981.

Who was Lyall Watson? Before we move on to the things that make the book unusual, we must ask: who was Lyall Watson? I recall being surprised on first learning of the existence of this book given that Watson was, and still is, best known for his 1973 Supernature: The Natural History of the Supernatural, a book on inexplicable phenomena and how they might be connected and explained. Supernature reads much like woo today, and it’s not surprising that Watson was regarded as embarrassingly credulous and even dishonest by some, and as refreshingly open-minded by others. I knew Watson for these reasons before discovering (by chance, in a bookshop… pre-internet days, kids) that he’d published a book on whales.

At left: Dr Lyall Watson. At right: 1973’s Supernature, Watson’s most famous book. Images: list of quotations of Lyall Watson (here), goodreads.com (here).

Caption: at left: Dr Lyall Watson. At right: 1973’s Supernature, Watson’s most famous book. Images: list of quotations of Lyall Watson (here), goodreads.com (here).

A look at the titles of his more than 20 published works reveals a remarkable and eclectic interest in all of natural history, in sport, culture and ritual (witness the 1989 Sumo: A Guide to Sumo Wrestling), in biology, anatomy and evolution, in the elements and physical geography, in the paranormal and spiritual, and in the human experience and everything about it. Many of us are interested in most or even all of these things, but scarcely any have the skill and knowledge that might allow us to write books on them. In keeping with his diverse interests and writing abilities, he was tremendously qualified, holding degrees in botany, zoology, ecology and anthropology. He even studied palaeontology under the great Raymond Dart. Watson completed his PhD on animal behaviour at the University of London under Desmond Morris, another scientist and author well known for a diverse skillset and ability to write engagingly about remarkable and controversial subjects. Unsurprisingly, Watson moved into the world of TV and also worked as a consultant for zoo and safari park design. Watson died in 2008 and there are some very good obituaries available online.

Anyway, back to the book. What makes it unusual?

Several of Ritchie’s whales, composited together (it might be obvious that I especially like beaked whales). Clockwise from upper left, we’re seeing Fraser’s dolphin Lagenodelphis hosei, Peale’s dolphin Lagenorhynchus australis, Strap-toothed whale …

Caption: several of Ritchie’s whales, composited together (it might be obvious that I especially like beaked whales). Clockwise from upper left, we’re seeing Fraser’s dolphin Lagenodelphis hosei, Peale’s dolphin Lagenorhynchus australis, Strap-toothed whale Mesoplodon layardii, Rough-toothed dolphin Steno bredanensis and Blainville’s beaked whale M. densirostris; Baird’s beaked whale Berardius bairdii is the big animal in the background. Images: Tom Ritchie/Watson 1981.

Whales of many hues. A key aspect of this book concerns its fantastic artwork. The whales look accurately proportioned and each illustration is nicely detailed. They’re not by Watson, but by artist Tom Ritchie. Watson states in a foreword how he and Ritchie travelled far – both north and south, he says – aboard the MS Lindblad Explorer in search of cetaceans. When describing the field sign, appearance and behaviour of cetaceans, he often describes things from the point of personal experience. Watson also states that he and Ritchie looked at numerous specimens in museum collections and also that they had access to new data never before published: the image of the Vaquita Phocoena sinus – named Gulf porpoise in the book (more on taxonomy in a minute) – “is taken from life and is the first ever printed which shows what the animal looks like” (Watson 1981, p. 8).

Ritchie’s Vaquita - at top - is apparently the first published full-body depiction of this animal’s life appearance. Below, a photo of a Vaquita in life. Extinction looms for this small cetacean. Images: Tom Ritchie/Watson 1981, Paula Olson/NOAA, in…

Caption: Ritchie’s Vaquita - at top - is apparently the first published full-body depiction of this animal’s life appearance. Below, a photo of a Vaquita in life. Extinction looms for this small cetacean. Images: Tom Ritchie/Watson 1981, Paula Olson/NOAA, in public domain (original here).

In view of all this, I find it fascinating that Ritchie’s cetaceans are often more boldly and brightly marked than those illustrated in other works, and depicted in hues that look surprising in view of more typical reconstructions (yes, it might be justifiable to term some depictions of living cetaceans reconstructions, since they’ve been cobbled together from diverse lines of evidence). The classic example is Stejneger’s beaked whale Mesoplodon stejnegeri (termed the Bering Sea beaked whale in the book). Photos and comments on this whale in living state show that it’s greyish brown, pale ventrally, and with off-white around the mouth and eyes. Ritchie’s version is warm brown dorsally, blue on its sides, white ventrally, and with a dark mask across the forehead and eyes (Watson 1981, p. 139). It’s an enhanced, technicolor version of the whale, and so different from other takes on this species that you’re left wondering how accurate it is. This sort of thing occurs throughout the book. The illustrations and wonderful and really attractive, but it’s difficult to be sure that they’re trustworthy.

Ritchie’s depiction of Stejneger’s beaked whale Mesoplodon stejnegeri. The hues and pattern depicted here are very different from other takes on the appearance of this animal. Image: Tom Ritchie/Watson 1981.

Caption: Ritchie’s depiction of Stejneger’s beaked whale Mesoplodon stejnegeri. The hues and pattern depicted here are very different from other takes on the appearance of this animal. Image: Tom Ritchie/Watson 1981.

I once wrote an April Fool’s article whereby a newly designed machine was said to have revealed the true life appearance of whales (it’s here at TetZoo ver 3). The imaginary multi-coloured whales devised for that spoof article were in part inspired …

Caption: I once wrote an April Fool’s article whereby a newly designed machine was said to have revealed the true life appearance of whales (it’s here at TetZoo ver 3). The imaginary multi-coloured whales devised for that spoof article were in part inspired by Tom Ritchie’s illustrations. Images: Gareth Monger and Darren Naish.

A heterodox phylogeny and taxonomy. A great strength of Watson (1981) is that it includes a fairly decent exposition on cetacean evolutionary history (now very dated of course) and copious discussion throughout of how anatomical characters group species together. What makes the book look odd today, however, is that Watson’s ideas are often heterodox and discordant with consensus views on these issues. We might expect no less of Watson given his other writings, but we might also wonder if the urge to shake things up a bit and promote new or minority opinions was a product of the time in which Watson was working (the late 1970s).

An early section in the book explains how the two great cetacean groups – mysticetes (baleen whales) and odontocetes (or toothed whales) – can’t definitely be said to share a recent common ancestor and might have emerged independently, and it’s even implied that this might also be true of ‘archaeocetes’, the archaic cetaceans otherwise regarded as the ancestors of mysticetes and odontocetes. Cetacean polyphyly is a weird idea in view of how many details mysticetes and odontocetes share to the exclusion of other mammals, but it would have seemed new and exciting during the 1970s given that it had come to the fore in papers of the mid and late 60s (Yablokov 1964, Van Valen 1968). Watson (1981) opted to support it. It isn’t taken seriously today, the anatomical, fossil and molecular evidence supporting cetacean monophyly being overwhelmingly good.

It gets better. Watson (1981) also opted to follow some (otherwise mostly ignored or forgotten) taxonomic proposals for delphinoids, and recognised a distinct Stenidae for ‘coastal dolphins’ (Steno, Sousa and Sotalia) and Globicephalidae for pilot and killer whales and their close kin. Those familiar with the technical literature on delphinoid evolution will know that both names originated elsewhere and have complex histories (which I must avoid discussing here), but their use in a field guide was unusual and heterodox given the tradition of including all of these animals within Delphinidae.

Watson (1981) wasn’t the only popular volume of the late 20th century to adopt some aspects of ‘non-traditional’ taxonomy. Anthony Martin et al.’s 1990 Whales and Dolphins also includes a globicephalid section (Martin 1990), which opens with this fa…

Caption: Watson (1981) wasn’t the only popular volume of the late 20th century to adopt some aspects of ‘non-traditional’ taxonomy. Anthony Martin et al.’s 1990 Whales and Dolphins also includes a globicephalid section (Martin 1990), which opens with this fantastic artwork (by Bruce Pearson). Image: Bruce Pearson/Martin 1990.

I should add that, in other respects, Watson (1981) seems conservative. Caperea is included within Balaenidae, the Kogia whales are included within Physeteridae (rather than their own Kogiidae; in this instance Watson states a preference to stick with consensus) and all river dolphins are lumped into Platanistidae, as was tradition at the time (though he noted that “There ought to perhaps be at least 3 separate families”, p. 148).

Watson’s Whales of the World includes various montage illustrations like this, which depict the field signs and characteristic markings of groups of species. The pictures look great. However, it has been argued that some of the details shown here ar…

Caption: Watson’s Whales of the World includes various montage illustrations like this, which depict the field signs and characteristic markings of groups of species. The pictures look great. However, it has been argued that some of the details shown here are not wholly reliable (read on). Images: Tom Ritchie/Watson 1981.

Smash the patronymy. On the subject of taxonomy – this time on common names rather than scientific ones – another bold move is the assertion that an overhaul is needed in naming conventions, and that biologists and naturalists should absolutely move away from the time-honoured tactic of naming animals after people. After all, calling a given animal – say – ‘Smith’s mouse’ tells you nothing at all about the mouse, does nothing to honour the remarkable features of said mouse, and is positively unhelpful should you see said mouse in the field and wish to remember its name. No, it should be the Epic blue mouse, or the Great spectacled forest mouse, Watson opined. I agree with this idea and also think that names should honour organisms. With this approach in mind, you won’t, then, find True’s beaked whale, Commerson’s dolphin or Bryde’s whale in Watson’s Whales of the World, but the Wonderful beaked whale, Piebald dolphin and Tropical whale, respectively (Watson 1981). Many new names of this sort are proposed in the book.

Close-up of Ritchie’s illustration of Shepherd’s beaked whale Tasmacetus shepherdi, one of my favourite living cetaceans. But it isn’t called Shepherd’s beaked whale in Watson (1981). Instead, it’s the Tasman whale. Image: Tom Ritchie/Watson 1981.

Caption: close-up of Ritchie’s illustration of Shepherd’s beaked whale Tasmacetus shepherdi, one of my favourite living cetaceans. But it isn’t called Shepherd’s beaked whale in Watson (1981). Instead, it’s the Tasman whale. Image: Tom Ritchie/Watson 1981.

However… language works best when we understand what other people are saying. When a word or name or turn of phrase is established and used throughout a community, it makes sense to stick with it, even if it’s misleading, technically inaccurate, or downright ‘wrong’. We can change it, but – I’d argue – we need to do so democratically, with input from as many relevant players as possible. I suppose a counter-argument is that someone has to get the ball rolling, and that proposing a new set of names in a book designed to function as a fieldguide is a good place to start.

Whatever the argument. Watson’s proposals didn’t win any accolade and his new names never became adopted by the cetological community. Maybe this was because he was an ‘outsider’ and lacked an established reputation as a whale expert or field biologist, but my main feeling is that most workers have wanted to stick with convention and continue to use the names that are otherwise entrenched.

My own whale illustrations - these were produced for various articles published back in the 1990s - were heavily inspired by those of Tom Ritchie. The originals of these illustrations appear to be lost today, so I have to draw them all anew for my i…

Caption: my own whale illustrations - these were produced for various articles published back in the 1990s - were heavily inspired by those of Tom Ritchie. The originals of these illustrations appear to be lost today, so I have to draw them all anew for my in-prep textbook. Image: Darren Naish.

The reception to Whales of the World. Having just noticed that Watson was seen as “an outsider”, it’s worth finishing this article by wondering how Whales of the World was received and perceived by specialists. Among whale researchers in general, the book was mostly ignored and generally regarded as problematic. Typical comments were provided by marine mammal specialist Niger Bonner (who wrote several excellent volumes on pinnipeds and cetaceans himself). Bonner noted that the book had noble aims but was marred by errors and erroneously gave the impression that many of the species were far better known than they really were (Bonner 1983). He criticised the maps, thought that the new naming system was arbitrary, confusing and annoying, and noted that the colours given to the animals in the artwork didn’t always match what was stated in the text (Bonner 1983).

So far as I can tell, these comments were and are typical, and what was – and remains – a popular and much-read book by amateurs and enthusiasts was never endorsed or recommended by those who know whales best.

Of all the popular and semi-technical books on cetaceans and other marine mammals, Watson (1981) remains one of the most interesting and attractive. This photo is from 2015 and I’ve acquired quite a few additional relevant volumes since. Image: Darr…

Caption: of all the popular and semi-technical books on cetaceans and other marine mammals, Watson (1981) remains one of the most interesting and attractive. This photo is from 2015 and I’ve acquired quite a few additional relevant volumes since. Image: Darren Naish.

I’m not a whale specialist, but I love the book, the caveat being – as should be obvious by now – that I love it for its weirdness and its design and artwork, not because I’ve ever found it an indispensable go-to work or a definitive take on the whales of the world. I’d say you should definitely get hold of it if you want a somewhat quirky, exciting take on the subject, or if you’re a completist or want to see Watson’s take on phylogeny, taxonomy and cetacean life appearance.

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Cetaceans have been covered at length on TetZoo before - mostly at ver 2 and ver 3 - but these articles are now all but useless since all their images have been removed (and/or they’re paywalled, thanks SciAm). Here are just a few of them…

Refs - -

Bonner, N. 1983. [Review of] Sea Guide to Whales of the World. Oryx 17, 49.

Martin, A. R. 1990. Whales and Dolphins. Salamander Books Ltd, London and New York.

Van Valen, L. 1968. Monophyly or diphyly in the origin of whales. Evolution 22, 37-41.

Watson, L. 1981. Whales of the World. Hutchinson, London.

Yablokov, A. V. 1964. Convergence or parallelism in the evolution of cetaceans. Paleontological Journal 1964, 97-106.