Disclaimer 2: The common names I use are all from the List
of Marine Mammal Species and Subspecies published by the Society for Marine
Mammalogy; these common names are generally not debated, though SMM writes: “common
names are arbitrary and change with time and place.” Nevertheless, common names
for individual species seem to be quite stable and meaningful to some degree;
it is the bigger labels – and common names of higher taxa – being dissected
here. You can view the list here.
I’d rather be talking about our new study on the large
extinct killer dolphin Ankylorhiza tiedemani, but after speaking with several
journalists about it I’m instead motivated to have a brief rant discussion
of the vernacular taxonomy of cetaceans. I get asked (or see) these sorts of questions/statements
frequently:
“So is it a
whale or is it a dolphin?”
“That’s a
whale tooth, not a dolphin tooth”
“The killer
whale is actually a dolphin”.
These words are used with such imagined precision and it
breaks my little heart every time because, well, they don’t mean a whole lot.
So many arguments are had over the identification of cetacean fossils, for
example, when in fact different completely defensible alternative definitions
exist and so many are comparing apples and oranges. And, I’ll note right now: I’ve
never heard an academic in mammalogy or paleontology ever get hung up on what
the definition of these mean, because we all A) tacitly acknowledge that they’re
not very well-defined and a bit meaningless in practice and B) use scientific
terms that have extremely precise meanings. So, a quick break down of the
terms.
Whale – from the Old English hwael, Old Norse hvalr,
Dutch & German wal, etc.
Definition according to Merriam Webster Dictionary: “any of various very large, aquatic, marine mammals (order Cetacea)
that have a torpedo-shaped body with a thick layer of blubber, paddle-shaped
forelimbs but no hind limbs, a horizontally flattened tail, and nostrils that
open externally at the top of the head”
Dolphin – from the Greek delphin,
Latin delphinus, old French dauphin.
Definition according to Merriam Webster Dictionary: “A)
any of various small marine toothed whales (family Delphinidae)
with the snout more or less elongated into a beak* and the neck vertebrae
partially fused. B) any of several related chiefly freshwater toothed whales
(as of the families Platanistidae and Iniidae)”
*beak, of course, is the cetological
term for the snout in a cetacean that protrudes beyond the melon – think of the
bottlenose in a bottlenose dolphin.
Porpoise – from the old French
porpais, translating to “pig fish” – originally from Latin porcus + piscus, porc
+ peis in French.
Definition according to Merriam
Webster Dictionary: “a blunt-snouted usually dark gray whale (Phocoena
phocoena) of the North Atlantic and North Pacific that
typically ranges from 5 to 6 feet (1.5 to 1.8 meters) in length.”
The definition of “whale” above perhaps
is the most broad, and characterizes dolphins and porpoises as well. The
definition of “dolphin” at first seems to only include the Delphinidae, but
then admits that there are some other dolphins – mostly riverine – that are
similar-ish in external body form that can be called dolphins. The definition
of porpoise offered by Merriam Webster highlights one species, the harbor
porpoise. Dolphins v. porpoises seems rather clear here: dolphins have beaks,
and porpoises do not. It is from these definitions that many accept a quasi-taxonomic
use of the words: dolphin means “Delphinidae” and porpoise means “Phocoenidae” –
the families containing the bottlenose dolphin and harbor porpoise
(respectively).
What about other definitions? I’ve read
elsewhere that a whale could be defined as any cetacean lacking a beak but
being larger than 9 feet in length, and that porpoises are any cetacean lacking
a beak smaller than that, and in many variants, possessing a triangular rather
than hooked (falcate) dorsal fin. This at first seems fine: phocoenids mostly
have triangular dorsal fins and none that live today possess a beak, clearly
differentiating them from delphinid species like the bottlenose dolphin.
However… what about the genus Cephalorhynchus? These are a fascinating
group of southern hemisphere delphinids with a “ring species” biogeographic
pattern of dispersal around the southern ocean – and none have a beak or a
falcate dorsal fin. Externally, they are so similar that if a stranded
individual were so decomposed that the teeth had fallen out and identification
was not possible based on the coloration, you would probably need to remove the
skull in order to tell which family it belonged to. On top of this, there has
been rampant use in the past of the word porpoise for delphinids with beaks
(spinner porpoise, bottlenose porpoise, etc.), as recently as the 1970s – in peer
reviewed papers by marine mammalogists like William Perrin!*
*I have also read this may be specific
to Hawaii, where local fisherman call all small cetaceans porpoises because
they instead use the term “dolphin” exclusively for the dolphinfish, which
really chaps my ass because there’s already a &#$ing local indigenous name
for that species – Mahi-mahi, that is now used more frequently than dolphinfish
outside of Hawaii.
Phylogenetic distribution of common names for cetaceans. Phylogeny from McGowen et al., 2011. Only the term porpoise, and only in its modern usage, appears to define an actual clade.
The distinction between “dolphin” and “porpoise”
is sadly, in my view, the clearest, and matters only become worse when you take
a closer look at “whale”. The first and biggest problem is that whale doesn’t
even attempt to mean anything biological: it includes both baleen whales and
large odontocetes, which, granted, are all cetaceans, but some whales (large
odontocetes) are obviously much, much more closely related to whatever dolphins
and porpoise are than others (e.g. mysticetes). There are two aspects to the
difference between whales and dolphins I’ve seen in most definitions or uses of
the words: 1) whales lack a beak and 2) whales are larger than dolphins. At
first glance, this works: baleen whales are all larger than dolphins, and sperm
whales, and killer whales are as well – and none of these have a beak. But,
there are, as before, exceptions.
The first exception are the beaked
whales: family Ziphiidae, all of which are larger than dolphins but… paradoxically
have beaks. One is even called the bottlenose whale! Another, Berardius,
is the second largest odontocete after the sperm whale Physeter, measuring
in at 11 meters or more – larger than minke whales, but smaller than the
largest bryde’s whales. Berardius, for the record, most certainly has a
beak.*
*I was lucky enough to participate in
a necropsy of a stranded female B. arnuxii on a frigid, foggy beach on
the southern tip of the south island of NZ during my Ph.D.
The second exception are some of the
smaller species of the “blackfish” – the Globicephalinae. These are a highly
derived group of darkly pigmented blunt-snouted delphinids with some suction
feeding adaptations. Despite being large bodied, darkly pigmented, and lacking
a beak, the killer whale is not actually a member of the Globicephalinae, and
is instead allied with a number of robust Tursiops-like fossils in the
Orcininae, the earliest diverging lineage within extant Delphinidae. Three of
these are the same size as most dolphins – 6-10 feet in length (2-3 meters) –
these are the melon-headed whale Peponocephala electra, pygmy killer
whale, Feresa attenuata, and the snubfin dolphins, Orcaella
spp. (Irrawaddy river dolphin, Australian snubfin dolphin). Peponocephala
(8-9 feet, 2.6-3 meters) and Feresa (6-7 feet, 2-2.3 meters) illustrate
that not all “whales” are large-bodied. Orcaella, the snubfin dolphins,
are small (6-9 feet, 2-3 meters), and called dolphins yet lack a beak! If your
brain wasn’t already close enough to melting out through your ears, let us also
consider the relatively large delphinid Grampus griseus – the Risso’s
Dolphin – which is a large globicephaline, smaller than a pilot whale or the
false killer whale, but much larger than the rest of the globicephalines. It is
large, and lacks a beak – externally it looks the same as a pilot whale aside
from coloration.
These terms are old with origins in the whaling industry, pre-dating modern biology.
Whale also is frequently used as an
umbrella term to refer to cetaceans in general, thereby tacitly including
dolphins and porpoises rather than always being distinguished from them. The
term “toothed whales” is the broadly accepted vernacular name for the
Odontoceti, and likewise, “baleen whales” for the Mysticeti. In older literature
(prior to World War II) you will find frequent references to the amusingly
named “whalebone whales”, and after cursory reading, you realize (just quickly
enough to avert a brain aneurysm, in my case) that all whalebone whales are
baleen whales. As it turns out, “whalebone” is an old whaling term for baleen,
which would be cut into strips and sold on shore to be used for the boning in
ladies’ corsets.
Another term you may hear is the “Great
whales” – which is defined by the International Whaling commission as the most
economically viable whales for the purposes of whaling – most baleen whales
(except for Eden’s and Omura’s whales), and the sperm whale. This is obviously
not a taxonomically informative moniker.
In summary, just looking at the
distribution of modern cetacean common names indicates that “whale”, “dolphin”,
and “porpoise” have no consistently used, biologically meaningful definitions.
They are informal terms, with multiple accepted definitions.
Testing the water: an informal twitter
poll
I started writing this post, and then realized as an afterthought,
that I should post a poll on twitter asking folks to vote on what they think a
particular cetacean was – the intention was to ask my twitter followers “should
the common name of this species have whale, dolphin, or porpoise in it?” not “is
this a cetacean?” whereby whale would have been correct every time. So, who
knows how useful the results are – regardless, I used the exact imprecise
language I’m railing against in the rest of this post, so confused results do
speak for themselves. It’s also worth noting that at present I have perhaps
5,000 followers give or take, and while many are scientists, most are not, and
even many scientists are not guaranteed to be familiar with obscure cetacean
species I’ve intentionally chosen to prove a point. [Admittedly, I have jumped the gun an hour or so, as the polls are not yet finished, but I don't expect them to change much. If there is any change I will update them accordingly].
The first poll had an unlabeled
photograph of a dolphin-like animal leaping from the water – without scale. Most
respondents (~49%) called this a dolphin (with the remaining votes split (25% each) between porpoise and whale) – but it is not a dolphin. This is a female Sowerby’s
beaked whale (Mesoplodon bidens) – difficult to tell, but the pectoral
fins are pretty far from the head. Answer: WHALE. [photo credit: azoreswhalewatch.com]
The second poll has a similar looking
animal leaping out of the water – and most (70%) responded correctly. This is a
spinner dolphin (Stenella longirostris) known for its acrobatic spinning
breaches. This was formerly frequently called the spinner porpoise, so you get
a free pass if you wrote porpoise. Answer: DOLPHIN. (or porpoise?) [photo credit: Getty Images TV via youtube]
The third poll is just a cute little face
poking out of the water – a face that is dark and lacks a beak. This one was
clearly confusing, as only a slight majority – 41% - identified it as a
porpoise, and about 30% said whale or dolphin. This is not a porpoise, however –
it is a Heaviside’s dolphin (Cephalorhynchus heavisidii), one of the smallest
dolphins, and a beakless one at that – in the porpoise-mimic genus Cephalorhynchus.
Answer: DOLPHIN. [photo credit: S. Elwen, Namibian Dolphin Project]
The fourth poll shows a sadly dead, strikingly
countershaded animal with an absurdly large dorsal fin, a white eye ring, and black
lipstick. Most respondents (~70%) correctly said this was a porpoise. This is
indeed the spectacled porpoise, Phocoena dioptrica. That’s actually my
Ph.D. adviser R.E. Fordyce (U. Otago, Geology dept.) on the left*, and Steve
Dawson (U. Otago, Marine Sciences dept.). Like the Heaviside’s dolphin above,
it does not have much of a beak. Answer: PORPOISE. [photo credit: 1 News, NZ]
*This beautiful porpoise stranded on the Otago Peninsula while I was in the Fordyce lab - but it was September 2014 and I was in the throes of Ph.D. writing, and desperately attempting to complete my Ph.D. thesis before January 10, my arbitrarily set deadline (which I made). I naturally had to decline the invitation, as necropsies followed by detailed dissections (this is R.E. Fordyce we're talking about; detail is everything) eat up several consecutive days.
*This beautiful porpoise stranded on the Otago Peninsula while I was in the Fordyce lab - but it was September 2014 and I was in the throes of Ph.D. writing, and desperately attempting to complete my Ph.D. thesis before January 10, my arbitrarily set deadline (which I made). I naturally had to decline the invitation, as necropsies followed by detailed dissections (this is R.E. Fordyce we're talking about; detail is everything) eat up several consecutive days.
The fifth poll, just for fun, was my illustration
of the recently re-named Ankylorhiza tiedemani – which I call a ‘killer
dolphin’ or ‘giant dolphin’. Respondents were pretty split between calling this
a whale (~43%) or a dolphin (~45%), with a few for porpoise. Answer: ???
What’s the moral of the story from these
polls? These terms are helpful in some cases but otherwise prove my point that
they do not reflect biological groups. Reliance upon these terms can lead to
confusion.
Fossil whales, dolphins, and porpoises
– a whaleontologist’s perspective
Paleontologists have a very laissez faire
approach, because we are constantly befuddled by strange cetaceans that further
blur the (admittedly very crookedly drawin) lines between these terms. Are four
legged Eocene semiaquatic archaeocetes - like Pakicetus - whales,
dolphins, or porpoises? They look rather close to a dog to be called any (but
are typically referred to as whales). What about some early sperm whales that had
relatively long rostra and may have had a beak, like the Miocene Zygophyseter?
If we stick to the strict Dolphin= Delphinidae, and Porpoise = Phocoenidae taxonomy
– what about dolphin-like phocoenids with beaks (e.g. Piscolithax), or extinct
delphinoids, like Kentriodon, that are not delphinids, but also small-bodied
with a beak? What about large kentriodontids the size of pilot whales, also
with a beak? What about our newly named Ankylorhiza – is it a whale or a
dolphin?
A somewhat outdated but nicely illustrated tree of modern and extinct cetaceans, highlighting the number of wholly extinct groups discussed by paleontologists without incident; modified from Barnes et al. (1985: Marine Mammal Science).
The strict dolphin v. porpoise dichotomy is
perhaps somewhat useful for modern cetaceans (delphinids including
porpoise-like species, and others labeled as whales, notwithstanding), but
completely falls apart thanks to a slew of extinct dolphin-like species that
are not delphinids. This is not universal, but from 15 years in the field, I
can say that whaleontologists do the following things: 1) whale broadly is used
by us to mean cetaceans, and all archaeocetes are generally referred to as “whales”.
2) Dolphin is broadly used for any small odontocete, including those with short
rostra (e.g. some fossil globicephalines, the extinct xenorophid Inermorostrum),
but also for larger species with beaks. Large dolphins, like Ankylorhiza
or the Miocene Hadrodelphis, are simply called large dolphins or giant
dolphins. Squalodon is called a shark toothed dolphin by most. In
general, with the exception of some modern species, we generally use the term
dolphin to indicate any extinct odontocete that is not a sperm whale or a
beaked whale. 3) Porpoise is always meant to refer to the family Phocoenidae.
4) River dolphin may be convenient for modern species, but most extinct
relatives of the four modern genera – Pontoporia, Inia, Lipotes,
and Platanista – were fully marine. River dolphins are three or four
separate clades representing a minimum of four riverine invasions.
One thing that seems obvious from my look
back is that the terms dolphin and whale are frequently used for many extinct
clades – but always with a family name in tow: e.g. eomysticetid whales;
kentriodontid dolphins, etc.
Coronodon havensteini, a baleen whale with teeth - a common source of confusion in our museum. Photo by me.
Toothed whales, or Echolocating whales
Now that we’ve clarified paleontological
usage of whale, dolphin, and porpoise, it’s time to take a closer look at the
two major groups, and terms that at first glance *appear* to be fine.
Odontocetes are the toothed whales, and Mysticetes are the baleen whales. But,
these vernacular names become problematic and confusing when communicating about
fossils to a lay audience. Some modern odontocetes don’t have any erupted teeth
or only have some tusks (e.g. narwhal, most beaked whales, risso’s dolphin),
and many extinct odontocetes similary had only tusks or were completely
toothless (Inermorostrum, Odobenocetops, Australodelphis, Vanbreenia,
Dolgopolis). This is not too much of a problem, to be honest.
Instead, the real problem is the toothed
mysticetes – baleen whales with teeth. Toothed baleen whales are the earliest
lineages of mysticetes that still had teeth – like Coronodon from South
Carolina, the Aetiocetidae (north Pacific flat-snouted teeth + baleen bearing
mysticetes), the Mammalodontidae (big-eyed Australasian toothed mysticetes),
and the Llanocetidae (southern hemisphere large bodied toothed mysticetes with
large gaps between teeth). We recently had a comment on a social media post about
Coronodon exclaiming “baleen whales don’t have teeth!”
While I have no idea how to rebrand “baleen
whales”, I have found a bit of a helpful compromise in the past few years:
rather than used the far more problematic term “toothed whales”, which relies
upon modern species and rapidly falls apart after cursory consideration of
fossils, I’ve started calling odontocetes “echolocating whales” since
discoveries like Cotylocara macei suggest the origin of echolocation to
be very, very early within the Odontoceti, quite possibly at the base of the
clade itself. A major exception to this is a paper that Dr. Rachel Racicot
published (I was a coauthor) on an Olympicetus-like dolphin from the
Oligocene of Washington state that apparently could not echolocate, thereby
implying that echolocation evolved twice within Odontoceti – or that this
dolphin lost the ability to echolocate. This probably doesn’t matter much, to
be honest, when you consider that virtually all modern and extinct odontocetes
have structures otherwise indicative of some facial structures associated with
sound production – and that it’s possible that our Olympicetus-like
dolphin may have lost the ability to echolocate. Such a case is clear with Odobenocetops,
which either had a tiny melon (O. leptodon) or probably did not
have one at all (O. peruvianus). To clarify: there are far fewer
odontocetes that cannot/could not echolocate than there are whales with teeth
that are not odontocetes. Therefore, I think the term “echolocating whales” is
probably a better common name than “toothed whales”.
So, what the hell is an orca? A whale
or a dolphin?
Now we’ve come full circle. Is an orca a
killer whale, or a killer dolphin? It’s both, really. Because it’s a member of
the family Delphinidae, it’s totally cool to call it a dolphin. If found as a
fossil, whaleontologists probably would’ve just called it a giant dolphin. At
the same time, whale doesn’t really have a specific biological meaning, so it’s
also just as fine to call it a whale. If you’re going to raise the point, you
may as well indicate that the Delphinidae are an ecomorphologically disparate clade
of cetaceans including dolphins, some that look and behave exactly like true
porpoises, and some that are called whales. So, don’t argue with
people because when you get down in the weeds about dolphin v. whale, none of
it makes any goddamn sense.
A possible compromise? A
whaleontologist’s glossary of vernacular terms
The semiformal vernacular nomenclature
used by paleocetologists suggests that a slightly more precise way of talking
about cetaceans is easy and can substantially clarify communication with the
public. Here is a brief synonymy list of terms I’ve heard used in paleocetology
and marine mammalogy (or used personally without precedent). It avoids for the
sake of useless repetition names including the family (e.g. xenorophid
dolphins). If you have more suggestions, please let me know in the comments and I'll consider adding them.
Whale = any cetacean, archaeocete or neocete
alike
Ancient whales**** = archaeocetes
Baleen whales = Mysticeti
Toothed baleen whales** = toothed Mysticeti
True baleen whales or baleen-bearing
whales** = Chaeomysticeti
Right whales = Balaenidae
Rorquals* = Balaenopteridae
Gray whales = Eschrichtiidae
Pygmy right whales = Neobalaenidae
Toothed whales or echolocating whales = Odontoceti
Dolphin**** = 1) for extant taxa: any
odontocete without whale or porpoise in the common name; 2) for extinct taxa:
any odontocete that’s not a phocoenid, physeteroid, ziphiid, or globicephaline
Shark-toothed dolphins**** = “Squalodontidae”
Spear-toothed dolphins*** = Waipatiidae
Swordfish dolphins*** =
Eurhinodelphinidae
Porpoises = Phocoenidae
Oceanic dolphin = Delphinidae
Blackfish**** = Globicephalinae + killer
whales
River dolphin**** = Platanista, Lipotes,
Inia, Pontoporia & extinct allies (marine or otherwise)
White whales = Monodontidae
Sperm whales = Physeteroidea
Walrus faced whales = Odobenocetops
*names used by mammalogists but are
probably useless for science communication
**extinct subdivisions/subtaxa of these
could benefit from better common names, thus far the family name is used
exclusively
***names I use that may or may not ever
catch on
****names that in no way refer to
monophyletic groups
Odobenocetops = whalerus
ReplyDeleteI approve.
ReplyDeleteWhat about limiting “whale” to the crown group and using “stem-whale” for “archaeocetes”?
ReplyDeleteThere is no faster way to lose the attention of average folks than crown/stem!
ReplyDeleteProto-whale? Calling e.g., Indohyus a "whale" seems like it'd be confusing.
ReplyDeleteIndohyus is a tricky one, but aside from a long snout and carnivory, there's not much that Pakicetus has that Indohyus does not. It generally has not been called a whale, though, from personal experience.
ReplyDelete