What big teeth you have…

One of Charles R. Knight’s paintings of Smilodon fatalis, this one menacing a giant sloth stuck in tar (off panel).

There are few fossil mammals that are as impressive the saber-toothed cat Smilodon fatalis, but despite it’s fearsome dentition some recent new reports have suggested it was more of a pussycat when it came to bite strength. This seems to be counter-intuitive; how could such an impressive animal be associated with the term “weak”? Part of it has to do with word choice, but the larger issue has to do with the fact that the bite of Smilodon wasn’t as strong as that of some other carnivores (extinct and extant), as well as dentition and feeding ecology. This issue goes far beyond just one genus or species, however, as Smilodon was only one of many genera that bore massive canines. In fact, huge “sabers” have evolved over-and-over again in the mammalian lineage (see this post and also this post for information about the cat-like ones), including the famous fangs of the machairodontine felids (saber-toothed cats) and their look-alike nimravid relatives.

Lateral, anterior, and dorsal views of the herbivore Uintatherium (Note the prominent canines). From Marsh, O.C. “The Fossil Mammals of the Order Dinocerata.” The American Naturalist, Vol. 7, No. 3. (Mar., 1873), pp. 146-153

Skull of another member of the Dinocerata; “Loxolophodon cornutus” (today known as Eobasileus cornutus). Again, note the prominent canine. From Cope, E.D. “The Amblypoda (Continued).” The American Naturalist, Vol. 19, No. 1. (Jan., 1885), pp. 40-55.

Although this post will primarily be concerned with the great “sabercats,” large, dagger-like canine teeth having been evolved multiple times by many different unrelated animals during the course of life on earth. In some herbivorous creatures, like the extinct Uintatherium and even in the extant Musk Deer, the fangs reflect sexual dimorphism and probably sexual selection, but the sharp teeth don’t seem to have a prominent function in mastication or processing of food. Likewise, large canine teeth are present in living baboons (Papio sp.), and the sexual dimorphism exhibited between the dental equipment of the males and the smaller canines of the females has long been noted (males often yawn to show off their canines, the size of their teeth being very intimidating indeed). Do the same considerations of sexual selection and dimorphism hold true for the saber-toothed cats, too? Unfortunately, fossil evidence does not always allow comparisons of the two sexes, but extant big cats and some death-trap sites have provided some information to work with. From Salesa, et al. (2006);

Among the Carnivora, sexual dimorphism is more marked in canine size than in other dental features or skull size, and these differences can be related to the breeding system. Species in which a male defends a group of females tend to be more dimorphic than those with monogamous pairs or groups of males and females. Felids are dimorphic animals, but mainly in reference to body size, with the mane of male lions being a unique example of morphological variation between sexes among the family.

This makes sense; if a male keeps a harem of females and has to defend it from other males, the species is more likely to exhibit sexual dimorphism than not. In cats, however, it seems to be more about body size (and possibly characters that wouldn’t fossilize in extinct species) than about tooth size (which would serve important other functions, so any sexual selection would be mitigated by natural selection), although we can’t be sure of this being that there are no living sabercats to study. Personally, I think there could be a sexual-selection component in some groups, but the saber-canine is so prominent in so many extinct felids and nimravids that it is extremely doubtful that all the lineages converged on similar tooth structures because of sexual selection/dimorphism, the functional advantage of larger teeth likely coming first. A lack of sexual dimorphism when considering morphology as a whole, however, may suggest a more solitary lifestyle where territories may or may not overlap are maintained and direct competition for females is not as fierce, especially since the females move through territories rather than living with a male. Such a strategy may have been employed by the late Micoene sabercat Paramachairodus ogygia. Salesa, et al. (2006), working with an assemblage made up of many of the more basal felids, have even been able to come up with a hypothesis about life history of the ancient animals based upon their finds in Spain;

[T]he probable territorial behaviour for Par. ogygia would be very similar to that of jaguars, in which males defend large, overlapping territories that include smaller territories of several females. This model is similar to that of the leopard, but in this species male territories never overlap, which could explain the different sexual dimorphism index of this species with respect to Par. ogygia and jaguar…

So, if Par. ogygia behaved more like jaguars and leopards than lions, the presence of juveniles in the trap would be highly improbable, as is the case. But in addition to the scarcity of juveniles, the sample from Batallones-1 has another interesting feature: it is mostly composed of young adults, that is, individuals with the complete permanent dentition, but without any trace of wear. These animals, which would have recently become independent of their mothers, would not as yet have had any territory, moving instead through the ranges of other adults and being more easily attracted by an easy meal, such as carrion. This age distribution therefore suggests that the sample of Par. ogygia trapped in Batallones-1 corresponds to that fraction of non-resident young individuals, both males and females, which were in a phase of dispersion. In the case of leopards, such individuals are more daring – or less cautious – than adults, and they have been seen crossing rivers in spate, whereas resident adults only cross at times of lower water. It has also been noticed that among these individuals, males are even more inclined to make these incursions than females, which remain longer with the mother, especially if there is good availability of food. If this pattern of dispersion behaviour applied to the young adults of Par. ogygia, it is likely that they were trapped in Batallones-1 more often than the resident adults.

Saber-Toothed Felid and Nimravid diversity (click for a larger image). From Emerson, S.B., and Radinsky, L. “Functional Analysis of Sabertooth Cranial Morphology.” Paleobiology, Vol. 6, No. 3. (Summer, 1980), pp. 295-312.

While the life histories of extinct mammalian carnivores are interesting in and of themselves, it is the teeth and terrifying bite of the sabercats that we are most concerned with here. Smilodon is the celebrity of saber-toothed cats, but the fossil record preserves a wide diversity of carnivores with large canine teeth, and even within the larger groupings there are even more subdivisions, the skulls of saber-toothed felids being widely variable. As discussed in the background material, nimravids are saber-tooth look-alikes that diverged from a common ancestral line earlier than the carnivores that would give rise to Smilodon, but the two lines are still closely related and have undergone parallel evolution. There is still some reshuffling of taxa going on and the true evolutionary history/affinities of many of the forms is still being worked out, but most forms you’re likely to see grouped together at a museum fall into either the nimravid or felid camps. The focus of this essay, however, will be on felids, and although they are often discussed along with their nimravid cousins the larger amount of work has been done on the felids and so we must leave the nimravids.

With the felids, then, there seem to be three kinds of sabercat that hint at differing predatory tactics, prey, and habitat. Indeed, evolution did not create carbon copies of the same creature, barring life from becoming adapted to varying circumstances; there is more variety than would be first assumed if we based all our research on the presence of prominent canines. Instead, there seem to be three “ways of being” a saber-toothed cat, as outlined by Martin, et al.;

Saber-toothed carnivores… have been divided into two groups: scimitar-toothed cats with shorter, coarsely serrated canines coupled with long legs for fast running, and dirk-toothed cats with more elongate, finely serrated canines coupled to short legs built for power rather than speed. In the Pleistocene of North America, as in Europe, the scimitar-cat was Homotherium; the North American dirk-tooth was Smilodon. We now describe a new sabercat from the Early Pleistocene of Florida [Xenosmilus], combining the scimitar-tooth canine with the short, massive limbs of a dirk-tooth predator. This presents a third way to construct a saber-toothed carnivore.

Xenosmilus hodsonae, Homotherium cf. crenatidens, and Homotherium serum. From Martin, L.D., Babiarz, J.P., Naples, V.L., and Hearst, J. “Three Ways To Be a Saber-Toothed Cat.” Naturwissenschaften, Vol. 87, No. 1 (Jan. 2000), pp. 41-44

As Martin notes, there appears to be a number of adaptational “trade offs” that sabercats in North America and Europe were subject to; fast-moving gracile forms had shorter sabers, but stouter and more powerful forms had the longer, more laterally flattened canine teeth. The “third way” that combined characters from both groups was exemplified by Xenosmilus (which Martin, et al. say would have seemed more like a bear than a cat, despite actual evolutionary relationships to the contrary). Still, leaving the overall structure of the body aside for a moment, the arrangement and sizing of the teeth of the different groups can be very telling. Martin, et al. again lay out what the usefulness of the differing tooth arrangements;

When biting, the long sabers of dirk-toothed cats may have cut parallel slits for some distance before the relatively smaller incisors could be applied. In scimitar-toothed cats the shorter canines and longer incisors worked more as a unit, first cutting parallel slits with the canines, immediately followed by the incisor arc removing the strip of flesh. Such a large open wound would have bled profusely, traumatizing the victim. If the incisors and canines acted in unison, the torsional forces on individual teeth would have been reduced, resulting in fewer restrictions on bite placement. In felids the size of the sagittal crest is directly proportional to the forces exerted by the temporalis musculature. Scimitar-toothed cats have a sagittal crest that is generally less pronounced than that in their dirk-toothed contemporaries. In a modification of the typical scimitar-tooth condition, the new cat from Florida exhibits both an elongated sagittal crest and an enlarged temporalis muscle that would have permitted a stronger bite.

While such a passage might not seem significant at first, it shows that there is more going on in a sabercat’s skull that is important to biting than just the size or shape of the canines. The placement of the incisors, for instance, seem to make a difference in biting strategy and force, dirk-toothed cats like Smilodon exhibiting a condition where the incisors are out forward of the canines. When this is taken into account, as well as the length of the canines, it seems that the canines would slash for quite some distance before the incisors could be used at all in comparison to the scimitar-toothed sabercats, the placement of the incisors in scimitar-tooths seemingly strengthening the biting teeth at the front of the jaw. The sagittal crests of these creatures should also be taken into account, such structures giving students of paleontology an indication of how carnivores (or herbivores, in the case of gorillas) have been adapted to achieve higher bite forces. Such ridges atop the skull for muscle attachment are not unique to sabercats, however, and there are some animals that have taken the structure to even greater extremes;

The extinct “bear dog” Amphicyon at the AMNH. Note the size of the sagittal crest, the reduction of the bony enclosure around the eyes, and the large holes on the side of the skull for jaw muscle attachment.

The extinct “saber-toothed” creodont Hyaenodon at the AMNH. Again, note the sagittal crest, reduction of bone enclosure around the eye, and the large canines.

The skull of the nimravid Hoplophoneus on display at the AMNH. Note the size of the canines and sagittal crest in comparison with Hyaenodon and Amphicyon.

The skull of Smilodon on display at the AMNH.

The skull of the marsupial predator Thylacoleo at the AMNH. Note the large openings on either side of the skull for the jaw muscles.

Ventral view of the skull of Thylacoleo. From E.D. Cope’s “The Tertiary Marsupialia” in The American Naturalist, Vol. 18, No. 7. (Jul., 1884), pp. 686-697.

Looking at the various groups, all show adaptations that increase the amount of available muscle attachment to achieve more powerful bites, modifying the skull in two ways. First, a sagittal crest (as already discussed) is often present to some degree, often being greater in omnivores or bone-crushing carnivores as they require greater forces to crack hard foods (although recent research by Wroe, et al. suggest that bone crushers like Spotted Hyena might not have the highest bite forces). Likewise, the holes between the skull and cheek bones are often enlarged or widened (the extreme of this group being Thylacoleo), the more muscle that can pass from lower jaw to skull being directly correlated to bite strength. What is interesting about sabercats, when considering these factors, is that they seem to be in the middle. They don’t exhibit adaptations of the skull to the extreme as in Amphicyon or Thylacoleo, but they still exhibit changes allowing for powerful bites (strong enough to kill and consume prey, at least). The trend is obvious and has not been missed by reseachers, and Emerson says the following about it;

With enlargement of upper canines, skulls of paleofelid, neofelid, marsupial and, as far as the record shows, creodont sabertooths were remodeled in similar ways. This evolutionary convergence in cranial morphology is not surprising, since most of the modifications relate to allowing increased gape while retaining bite strength at the carnassial. Those are factors essential for all sabertooths, and the possible ways to achieve them, starting from a generalized mammalian cranial morphology, are limited…

Why did sabertooth specializations evolve so many times? Their multiple evolution, plus the fact that several species of sabertoothed felids existed for most of the history of the family (from about 35 Myr to about 15,000 yr BP) suggest that sabertooth canines provided an effective alternative to the modern carnivore mode of killing prey

The skull of the saber-toothed cat Megantereon. Like in Smilodon, not how the incisors jut out (as well as the overly large nasal opening in this genus).

The basic mechanics of the skull just discussed gives researchers clues as to how sabercats could have killed their prey, but reconstructing ancient predator/prey interactions with no exact modern equivalent is difficult. Indeed, debate has gone on for years as to how sabercats used their teeth to bring down prey (see Simpson’s paper), either by stabbing, cutting, slicing, or even (as silly as it may seem) by crushing. What does seem apparent today, however, is that the canines of the sabercats were relatively delicate, and it would be unwise to fully sink them into a struggling animal as they may easily be broken off. Even if such an attempt to deeply puncture a prey item was not undertaken, biting full-force into bone could have also easily damaged teeth (or even broken them off), making it unlikely that sabercats jumped onto the back of their prey and tried to sink their teeth into the back of the prey’s skull like some modern cats. Recent research has even shown that the skull of Smilodon was ill-suited to handle stresses associated with struggling prey when compared to the skull of a lion, and I wonder how often individual Smilodon perished because of stresses associated with taking down prey if the victim was not brought down and killed quickly. Indeed, it seems that the long teeth were better suited to slicing soft flesh, i.e. cutting open the belly of prey or slicing open the throat, rather than piercing rough hides and ramming through bone.

Skulls (mandibles not pictured) of 4 “saber-toothed” mammals from “The Function of Saber-Like Canines in Carnivorous Mammals” by G.G. Simpson, American Museum Novitiates, August 4, 1941. Pictured are A) Machairodus (felid), B) Hoplophoneus (nimravid), C) Smilodon (felid), and D) Thylacosmilus (marsupial).

As just discussed in terms of tooth and skull stressed, many factors of life history, behavior, and morphology of extant big cats and sabercats might be similar, but the massive canines of the extinct group seem to infer a different killing strategy, and there is no reason to assume that they were like modern big cats in every respect. Salesa, et al. sums it up this way;

Extant felids kill small animals by biting on the nape or directly on the skull, using their rounded-section canines, but if any sabre-toothed cat tried to do this they would have risked breaking the laterally flattened upper canines. For this reason, it is more probable that they developed some behavioural mechanism to minimize that risk, such as ignoring prey below a given size. It is likely that machairodontines developed this ethological trait early in their evolution, and so narrowed their prey size range in comparison with that of felines, which hunt both large and small animals. This high specialization has been pointed out as one of the possible reasons for the gradual decline and final extinction of the sabre-toothed cats in the Pleistocene… The development of this strategy was probably the key reason for the sabre-tooted cats becoming the dominant predators in the land mammal faunas from the Late Miocene to Late Pleistocene.

It might not immediately make sense that felids with fragile teeth would specialize in eating large prey, but that is whale the fossil evidence (as we currently understand it) infers. While the smallest prey would pose no problems (outside of not being a fully satisfying meal), but medium sized prey with smaller areas of soft flesh (like the stomach and neck) would potentially be more dangerous and a more exact bite would be needed to prevent damage to the teeth and skull. Hence, it seems that the slashing and ripping of soft tissue in larger animals was the main method of killing prey (after it had been brought down or slowed by blood loss), taking hypercarnivory to an even more specialized extent.

An Amur Leopard yawns. Note the relatively small (but still fearsome) canines of the upper and lower jaw.

What, then, of a smaller living cat, the Clouded Leopard (Neofelis nebulosa and N. diardii), which has been heralded as a modern analog of sabercats? As Christiansen notes, Clouded Leopards are a bit bizarre, and it is incorrect to call them “small” big cats or modern sabercats, the genus showing a number of convergences with extinct forms while remaining distinct from the famed genus Panthera;

The skull morphology of the clouded leopard sets it apart from other extant felids, and in a number of respects it approaches the morphology of primitive sabertooths. This indicates convergence of several characters in machairodontine felids and the clouded leopard, mainly as adaptations for attaining a large gape. This raises doubts about the characters hitherto considered as distinguishing sabertoothed from nonsabertoothed predators…

Clearly, Neofelis and the sabertooths independently evolved a suite of the same specializations for the same overall purpose of attaining a large gape, a prerequisite for efficient jaw mechanics with large canines, but the reasons for evolving these characters need not have been similar. Based on analyses of lower jaw bending moments and inferred resistance to mechanical loadings, Therrien (2005) suggested that Neofelis could be at the beginning of a new sabertooth radiation. Such claims are difficult to test, however, since the extant sister taxon to Neofelis (Panthera) shares none of its sabertoothed characters, and the fossil record provides no clues of felids closer to Neofelis than Panthera. At present, however, there is little evidence to suggest that Neofelis can be regarded as an “extant sabertooth,” although it clearly shares a number of characters with them that are absent in other extant felids. On the other hand, it cannot be regarded as simply an intermediate between large and small felids, as normally assumed. The presence to some extent of characters normally ascribed to sabertooths in Neofelis raises doubts about their functional and evolutionary significance in primitive machairodonts such as Nimravides or Paramachairodus, hitherto the only reasonably well-known primitive machairodont. Such animals need not have shared the presumed functional skull morphology of later, more derived sabertooths and are perhaps not to be regarded as “sabertoothed” at all, if by sabertoothed is implied animals functionally significantly different from extant felids.

Again, this shows a convergence of functional morphology despite existing evolutionary relationships, many felids being adapted in similar ways. As stated previously, the large canines of saber-toothed predators required the animals to open their jaws wide but also narrowed their predatory niche to some extent. Likewise, various tests seem to show that the bite of sabercats like Smilodon was “weak,” with news reports often relating that the terrible felids were more like big housecats when compared to living big cats. This is a mistake (and it would be a grave one for anyone ever to cross a sabercat), born of a lack of recognition that bite forces exist on a continuum and are related to a number of factors and cannot simply be deemed “weak” or “strong” without further comment. Christiansen relates the bite force of Smilodon as such;

[A]lthough large sabertooths such as Smilodon and Homotherium had weaker bite forces than lions or tigers, their bite forces were broadly comparable to those of jaguars and large leopards, and, thus, cannot be claimed to have been “weak”. Lower bite forces at any given body size were probably evolutionarily possible owing to a marked contribution from the upper cervical musculature to the killing bite, which… was absent in Neofelis and primitive machairodonts such as Paramachairodus. Thus, bite force analysis may constitute a hitherto overlooked parameter in evaluating whether or not primitive machairodonts such as Paramachairodus or Nimravides really did employ a canine shear bite with a marked contribution from the cervical muscles to subdue prey, or killed in a manner similar to extant felids, which requires a stronger killing bite…

In many Plio-Pleistocene communities predator competition was more severe than today, and a sabertooth killing mode could be a way of ensuring faster kill rates, since a throat shear-bite most likely would kill prey faster than a throttling throat bite, common in extant pantherines. In lions, it can take up to 13 minutes to kill large prey, and in such cases the prey is frequently killed by disemboweling by other pride members. In the cheetah a suffocation bite can take even longer to kill prey. Carcass theft and feeding competition is very common among extant large, sympatric predators, and a faster kill mode could be a way of reducing the risk of carcass theft from competing predators. In many large predators with sympatric competitors, rapid consumption can be a way of reducing the risk of carcass theft, and this would most likely have been accentuated in past ecosystems with more intense large predator competition. Accordingly, the morphology and behavior of extant predators need not reflect the circumstances to which they became adapted when they evolved. More intense competition could accelerate the evolution of a sabertooth morphology…

This passage reflects the problems with reconstructing bite forces and predation techniques of extinct creatures; more is involved than just the opening and closing of the jaw. The neck muscles of many sabercats (except in some of the more basal members, as noted) likely contributed to the strength of the bite in a way that’s not directly testable today. Likewise, the killing technique of sabercats might not have required a bite as strong as a modern-day tiger, as in a land filled with other predators, it might simply take too long to try and suffocate a prey animal or bite through the back of their skull. Disemboweling or tearing out the throat of the prey item, by contrast, is a much quicker way to do large amounts of damage but it seems that it would require teamwork, solitary extant big cats often opting for a killing neck bite when the prey has been brought down. Even if this is eventually shown to be incorrect, it should be remembered that bite strength is not everything; despite its large size, the Great White Shark (Carcharadon carcharias) has a relatively weak bite, but it makes up for it with heavily serrated teeth, force of impact when attacking prey, and side-to-side head shaking to saw through its food. Crocodilians, by contrast, have very strong bite forces but they don’t saw through prey or chew, the emphasis being holding on to struggling prey and drowning it before ripping it apart. Such considerations bring us to another point mentioned above in our discussion of scimitar-tooths vs. dirk tooths in that the famous dirk-toothed cats like Smilodon were more powerfully built, seemingly focusing on bringing a large animal down to the ground and then delivering devastating bites once the stomach and neck were exposed (a process that would be made easier by groups working together, as seen in modern examples like lions bringing down giraffes or elephants).

A group of lions brings down a giraffe.

A group of lions brings down an elephant.

A new paper, just out in PNAS, does take the powerful neck muscles of Smilodon into account, however, and the information from the new models appear to corraborate the modern understanding of a felid that captured and killed prey in a way quite different from Panthera. From McHenry, et al.;

Our results demonstrate that bite force driven by jaw muscles was relatively weak in S. fatalis, one-third that of a lion (Panthera leo) of comparable size, and its skull was poorly optimized to resist the extrinsic loadings generated by struggling prey. Its skull is better optimized for bites on restrained prey where the bite is augmented by force from the cervical musculature. We conclude that prey were brought to ground and restrained before a killing bite, driven in large part by powerful cervical musculature. Because large prey is easier to restrain if its head is secured, the killing bite was most likely directed to the neck. We suggest that the more powerful jaw muscles of P. leo may be required for extended, asphyxiating bites and that the relatively low bite forces in S. fatalis might reflect its ability to kill large prey more quickly, avoiding the need for prolonged bites.

Hunting isn’t the only aspect of sabercat predation that seems to have differed from modern carnivores; they way they ate (and what they ate) is somewhat at variance with modern forms, as well. As is apparent at this point, the contact of the canines with bones would have been avoided, and it seems that the hard parts of the skeleton would have been avoided when a sabercat was consuming it. This could differ among different groups (perhaps some of the shorter-toothed forms not being so finicky about bone), but research into microwear patterns on teeth of Smilodon don’t seem to match with wear patterns of any living carnivores, suggesting a different dietary preference. It could be hypothesized, then, that creatures like Smilodon primarily consumed the soft parts of the carcass or what could be removed without too much damage to the teeth, and it should be remembered that living big cats often do not eat every part of the skeleton. Some, like cougars, have favored parts that they eat but end up leaving as much as 40% of the carcass behind. Other predators, especially bone-crushing ones, could take advantage of the leftovers, although the felids might have had to eat quickly as some of their osteophagus competitors may not have been patient (and, in fact, lions and hyenas often fight over kills and steal them from each other today).

Given all the prior considerations, it now seems that sabercats specialized in bringing down relatively large prey down quickly (some likely working in groups to do so), killing the victims by slashing open their stomachs or slicing through the blood vessels of the neck. This would be a much messier, but quicker, method than employed by living big cats, although the limitation of food sources likely caused in the eventual downfall of sabercats. Hypercarnivory can be a dangerous adaptive path to go down, and cats are clearly the most meat-dependant of the Carnivora, but it seems that extinct forms took their dental and dietary specialization above and beyond what is seen today. The price paid for such adaptations ended up being extinction, but given how many times they have shown up in the history of life on this planet, someday there may again be a saber-toothed predator stalking the shadows.

References;

Anyonge, W. “Microwear on Canines and Killing Behavior in Large Carnivores: Saber Function in Smilodon fatalis” Journal of Mammalogy, Vol. 77, No. 4 (Nov., 1996), pp. 1059-1067

Christiansen, P. “Canine morphology in the larger Felidae: implications for feeding ecology.” Biological Journal of the Linnean Society. Vol. 91, No. 4 (Aug., 2007), pp. 573-592

Christiansen, P. “Sabertooth characters in the clouded leopard (Neofelis nebulosa Griffiths 1821).” Journal of Morphology, Vol. 267, No. 10 (Jul., 2006), pp. 1186 – 1198

Christiansen, P. and Wroe, S. “Bite Forces and Evolutionary Adaptations to Feeding Ecology in Carnivores.” Ecology, Vol. 88, No. 2 (Feb., 2007), pp. 347–358

Cope, E.D. “The Amblypoda (Continued).” The American Naturalist, Vol. 19, No. 1. (Jan., 1885), pp. 40-55.

Cope, E.D. “The Tertiary Marsupialia.” The American Naturalist, Vol. 18, No. 7. (Jul., 1884), pp. 686-697.

Emerson, S.B., and Radinsky, L. “Functional Analysis of Sabertooth Cranial Morphology.” Paleobiology, Vol. 6, No. 3. (Summer, 1980), pp. 295-312.

Leutenegger, W., and Kelly, J.T. “Relationship of sexual dimorphism in canine size and body size to social, behavioral, and ecological correlates in anthropoid primates.” Primates, Vol. 18, No. 1 (Jan., 1977), pp. 117-136

Lucas, P.W., Corlett, R.T., and Luke, D.A. “Sexual dimorphism of tooth size in anthropoids.” Human Evolution Vol. 1, No. 1 (Feb., 1986), pp. 23-39

Marsh, O.C. “The Fossil Mammals of the Order Dinocerata.” The American Naturalist, Vol. 7, No. 3. (Mar., 1873), pp. 146-153

Martin, L.D., Babiarz, J.P., Naples, V.L., and Hearst, J. “Three Ways To Be a Saber-Toothed Cat.” Naturwissenschaften, Vol. 87, No. 1 (Jan. 2000), pp. 41-44

McHenry, C.R., et al. “Supermodeled sabercat, predatory behavior in Smilodon fatalis revealed by high-resolution 3D computer simulation.” PNAS, Published online before print October 2, 2007

Salesa, M.J., et al. “Aspects of the functional morphology in the cranial and cervical skeleton of the sabre-toothed cat Paramachairodus ogygia (Kaup, 1832) (Felidae, Machairodontinae) from the Late Miocene of Spain: implications for the origins of the machairodont killing bite.” Zoological Journal of the Linnean Society, Vol. 144, No. 3, (Jul., 2005) pp. 363-377

Salesa, M.J., et al. “Inferred behaviour and ecology of the primitive sabre-toothed cat Paramachairodus ogygia (Felidae, Machairodontinae) from the Late Miocene of Spain” Journal of Zoology, Vol. 268, No. 3 (Mar., 2006), pp. 243-254

Simpson, G.G. “The Function of Saber-Like Canines in Carnivorous Mammals.” American Museum Novitiates, August 4, 1941

Therrian, F. “Mandibular force profiles of extant carnivorans and implications for the feeding behaviour of extinct predators.” Journal of Zoology, Vol. 276, No. 3 (Nov., 2005), pp. 249-270

Therrian, F. “Feeding behaviour and bite force of sabretoothed predators.” Zoological Journal of the Linnean Society, Vol. 145, No. 3 (Nov., 2005), pp. 393-426

Van Valkenburgh, B., and Molnar, R.E. “Dinosaurian and mammalian predators compared.” Paleobiology, Vol. 28, No. 4 (Dec., 2002), pp. 527–543

Walker, Alan. “Mechanisms of honing in the male baboon canine.” American Journal of Physical Anthropology, Vol. 65, No. 1 (?, 1984), pp. 47 – 60

Wroe, S., McHenry, C., and Thomason, Jeffery. “Bite club: comparative bite force in big biting mammals and the prediction of predatory behaviour in fossil taxa.” Proceedings of the Royal Society B, Vol. 272, No. 1563 (Mar., 2005), pp. 619-625

The Chimpanzees of Mt. Assirik

When chimpanzees (Pan troglodytes) appear in documentaries they are often shown inhabiting relatively dense tropical forest, their lives taking place within the green refuge of the forests. As with any other species that is spread over a considerable distance, however, different populations of chimpanzees have different habits, and one of the most remarkable populations are those around Mt. Assirik. Located in the southeastern part of the Parc National du Niokolo-Koba in Senegal, the chimpanzees in this area have to deal with a local ecology that is drier and more open than some of their relatives elsewhere, and their behavioral adaptations to the environment is of great interest to those study human origins.

The Mt. Assirik study area is remarkable in that 55% of the habitat is open grassland, only about 37% being woodland of varying density and only 3% being more dense forest (the remaining area being made up of bamboo forest and isolated trees). Such open spaces allow some of the major Carnivora of Africa to live in close proximity to the chimpanzees; Lions (Panthera leo), Leopards (Panthera pardus), Wild Dogs (Lycaon pictus), and Spotted Hyenas (Crocuta crocuta) are all frequently seen in the area. As if having so many predators at their doorstep were not enough, the Mt. Assirik area seems to have fluctuations of food that aren’t correlated with seasonal changes, and in the dry season water is the most prized of any resource. The apes are not entirely helpless in the face of such pressures, however, and they’ve been behaviorally adapted in some very interesting ways.

Given a choice, the Mt. Assirik chimpanzees prefer to spend their time in the denser areas of forest, but shifting food resources sometimes require them to move across large expanses of open grassland in order to find nourishment. Wandering out onto the open plains alone is so dangerous as to nearly be suicidal, and the apes form large mixed groups when they have to move across the plains. During this time they are at their most vulnerable, especially since they would be unlikely to outrun any of the major predators (especially those that hunt in packs), and they are extremely alert when undertaking such a journey. What is perhaps most striking of all, hearkening back to Raymond Dart’s “Savanna Hypothesis,” is the fact that the chimpanzees sometimes stand up to get a better look at their surroundings, potentially spotting predators before they get too close, although such an observation should not be taken as a sweeping vindication of Dart’s ideas of human evolution.

The presence of just one tree or a few trees spaced far apart doesn’t help the chimpanzees much either; mothers with children and individuals spent much less time in the sparser woodland areas than in the forest, mixed groups seemingly having to issues with the woodlands. Why should this be so? Well, leopards can climb trees (and often do so to stash their kills), as well as lions, and so simply climbing a tree does not equal escape. Lone chimpanzees are far more comfortable in a habitat where they can climb a tree and move through the canopy out of reach of their assailants, something that is not possible in woodlands. The predators may also have another effect on the diet of the chimpanzees; the Mt. Assirik chimps do not seem to eat young ungulates or monkeys, although such behaviors have been made famous where it has been observed (i.e. Gombe). This may be due to some competition, but it may also be due to the restricted forested habitat and the fact that chimpanzees would have to enter the habitat of the carnivores in order to capture young ungulates, predators being likely to quickly learn about any kills that had been made.

Indeed, the Mt. Assirik population is remarkable in that it often moves long distances in order to obtain food as it becomes available, relying on numbers and vigilance to protect itself from predators when it’s habitat only offers a few isolated islands of relief. Although humans did not evolve from modern chimpanzees, this population may give researchers some idea of the behavior patterns of our ancestors when faced with similar constraints when forests became sparser and the plains were filled with predators. Such social behavior is not the only thing that makes the Mt. Assirik chimpanzees stand out, however; they also make use of Baobab trees in a very interesting way.

By now many people are familiar with the ability of chimpanzees to use a piece of wood as a hammer to break a nut placed upon an “anvil” of rock or tree root; such footage has been shown in television programs again and again. Such behavior did not come out of nowhere, however, and the way Mt. Assirik chimpanzees open nuts may represent a stage of tool use that precedes the hammer-and-anvil technology. While it had been disputed for some time whether the Mt. Assirik population used hammers and anvils or just anvils, recent studies have shown that they are cracking open the hard nuts of the tree on branches and not using a hammer. While we might think of an “anvil” as something that can only be used in conjunction with a hammer, mechanically this isn’t necessarily so, and the Mt. Assirik chimpanzees bang the hard nuts they collect on the branches of the tree (therefore staying aloft, not coming down to use stones or the roots of the tree), the tree itself being the anvil.

Given the basal usage of anvils by the Mt. Assirik chimps and the use of hammers and anvils elsewhere, it becomes possible to hypothesize about the evolution of stone tool use in our own ancestors. The starting point was likely similar to what is exhibited by the Mt. Assirik chimpanzees, banging hard nuts on trees or rocks in order to open them (thus preventing damage to the teeth, if it even would be possible to open the nuts using their jaws). The next step would be adding a hammer, possibly wooden (as seen in some groups today) or possibly stone. At this stage any combination of wood or stone hammers and anvils could be used, but tool use would probably not progress until a population was using stone hammers and stone anvils to open foods. In such a scenario, the apes would sometimes miss their targets and flake off bits of stone, an accident that would shape the tools. When a certain cognitive leap was made, the apes could then move from accidentally flaking their tools to doing it intentionally to truly be making tools rather than making use of naturally occurring bits of wood and stone. The reality of the situation may be forever lost to us, ancient tool use before the knapping of stone became prevalent being notoriously hard to discern, but such a line of behavioral descent is not unreasonable and seems to allow further development merely by chance combinations of naturally occurring resources.

Such a discussion is only a brief sketch based upon what I have only recently learned myself, but I hope that it has been at least somewhat informative. Different populations of chimpanzees show different behaviors and live in differing ecologies, and it would be a mistake to assume what the famous Gombe chimpanzees are doing holds true for all the other populations. Another population that I soon intend to write about spends time in caves, probes trees for bush babies, and may even have the beginnings of a fire culture; others do not show the same exact behaviors, but they have their own cultures and reactions to the local ecology. While we should be careful in analyzing the living populations of chimpanzees and their perceived similarities to humans, it would be foolish to think that they can tell us nothing of our own past, and if very well may be that some of traits (behavioral or otherwise) they now exhibit were present in our own lineage, vignettes of evolutionary history being replayed with different actors in our own time.

A peek at my homework

Here’s the summary that I’ll be giving today in my Topics of African Prehistory course pertaining to the assigned reading Wrangham, R. 1987. “The Significance of African Apes for Reconstructing Human Social Evolution.” In Warren G. Kinzey (Ed.) The evolution of Human Behavior: Primate Models. It’s long by summary standards, but when have I been known to be succinct? In fact, I would have loved to make this even longer, but I don’t want to talk my classmates to death.

Summary

“Life is a copiously branching bush, continually pruned by the grim reaper of extinction, not a ladder of predictable progress. Most people may know this as a phrase to be uttered, but not as a concept brought into the deep interior of understanding. Hence we continually make errors inspired by unconscious allegiance to the ladder of progress, even when we explicitly deny such a superannuated view of life.” – Stephen Jay Gould, Wonderful Life, 1989

On June 30, 1860, “Darwin’s Bulldog” T.H. Huxley met Bishop “Soapy Sam” Wilberfoce in a debate on one of the most hotly contested topics ever to be put before mankind: are we evolved, or are we divine creations? While no one is quite certain as to the outcome of the debate, it is perhaps one of the most celebrated events in the history of the evolution idea, for when the Bishop rhetorically asked whether it was through his grandmother or grandfather that he was descended from a monkey, Huxley delivered this devastating rejoinder; “If then the question is put to me would I rather have a miserable ape for a grandfather or a man highly endowed by nature and possessing great means and influence and yet who employs those faculties and that influence for the mere purpose of introducing ridicule into a grave scientific discussion – I unhesitatingly affirm my preference for the ape.” Such wit did not halt the debate then and there, but as Huxley’s work Man’s Place in Nature, Darwin’s The Descent of Man, and various cartoons from Punch at the time make clear, it could no longer be denied that Homo sapiens had very close relations to the living gibbons, orangutans, gorillas, and chimpanzees, their lives providing insight into our own.

The relationship between men and apes is now taken as a “given” (and rightly so), but the question of just what living apes can tell us about our past must be asked. While the fossil record seemingly refused to give up hominid remains for some time, there is today a greater diversity of fossil hominids now known than in Huxley’s time, and what we know about living apes must be reconciled with these discoveries if we’re to accurately portray what our ancestors (even our common ancestors) may have been like. Indeed, we should not forget that our own species did not evolve from chimpanzees or gorillas but rather shared common ancestry with them in the past, and they have been evolving since the time of their separation just as we have. As Richard Wrangham rightly criticizes the approach of trying to crown a living species as the archetype for our ancestor, noting “The ideas these models generate are plausible and even thought provoking, but their value is limited by their initial assumption: they assume that the social organization of human ancestors was similar to that of living species.”

Given this potential pitfall, Wrangham suggests a behavioral sort of cladistics, surveying the social behavior of extant gorillas, chimpanzees, and bonobos in order to find the presence (or absence) of shared social behaviors. If certain behaviors exist within all the groups mentioned, then there would be reason to believe (at least in terms of parsimony) that such behaviors were inherited from a common ancestor rather than evolved multiple times. Concerning the closed or semi-closed social groups detailed in II A 2. (Grouping Patterns) of the outline, it appears that humans, chimpanzees, and gorillas all have closed or semi-closed social groups, making the behavior a shared trait that may have been shared by the common ancestor of all the groups. On the other hand, however, we have the data presented in II B 4. (Male-Male Interactions) where the variety of interactions precludes us from being able to tell what sort of behavior pattern our common ancestor exhibited in terms of male interactions.

Now that we understand the application of Wrangham’s methodology to living primates, we should consider the overall strengths and weaknesses it may have. One of its strengths may be the ability to recognize the possession of common behavior in the apes despite different ecologies. If humans, chimpanzees, bonobos, and gorillas all share certain behavioral characteristics despite living in different habitats or inhabiting different niches, the overall case is stronger for that trait being inherited from a common ancestor rather than convergent evolution. Convergent evolution can be problematic, however, as it sometimes seems to defy parsimonious explanations. Perhaps the common ancestor did not exhibit the behaviors now expressed as much carry a capacity for them through variations in populations (being that it is populations that speciate and change, not an entire species as a whole), and being that we are dealing with behavior and not morphology in this case, it might be easy to accentuate some similarities/differences while hiding others. For example, if we undertake cladistics in the traditional sense, let’s say describing a skull, the process is relatively straightforward; either a structure or trait is present or it is not. Behavior, however, can be more variable, and even in Wrangham’s description of Group Patterns there can be seen some potential for disagreement. Indeed, is there are large significance between a closed group and a semi-closed group? Again, given that we’re talking about behavior and not a morphological trait that is usually clearly present or absent, researchers would do well to be mindful of how they delineate what they consider significant behaviors and how they are measured in terms of this method.

Wrangham does not hold his method up as the one and only answer, however, and he concedes that it is more of a “quick” and “weak” starting point to determine possible similarities rather than a way to obtain ultimate answers. In fact, as he notes in the introductory paragraphs, the study of behavioral ecology weighs heavily on the issues herein discussed, although it is a discipline still in development. Even beyond modern ecology, paleoecology will have a very significant role to play in determining what our ancestors were like, especially because habitat does much to shape the bodies and behaviors of organisms through evolution. Indeed, living species can give us valuable insights into our past, but if the information gleaned is determined to be the product of convergence or is found not to be consonant with the data from the fossil record, developing understanding will have to accommodate such discoveries. Ultimately, the discovery and determination of our common ancestors with chimpanzees and gorillas will weigh heavily on this issue, but at the present time the information is overall insufficient and (as ever) there are more questions than answers. For the present, however, the behavior of living African apes can provide a sufficient framework for comparison, and Wrangham’s methodology provides a quick way to spot potential similarities that can then be checked through the study of ecology and the fossil record.

There is a grandeur in this view of life…

A female gorilla at the Bronx Zoo.

It is often all-too-easy to forget about the wonder that is in nature when one becomes embroiled in the culture war surrounding evolution and creationism. The battles are fought in public classrooms, sundry media outlets, and (perhaps most of all) the internet, but those who do recognize the intricacy and beauty of evolution should not forget to step back every once in a while and look at what so-inspired Darwin in the first place. Nature offers up more treasures and wonders than I could ever fully appreciate during my short tenure on this planet, and without this sense of unity and amazement science can quickly turn into a rather dry and forbidding set of mental exercises.

Aldo Leopold recognized this problem all too well. In his essay “Song of the Gavilan”, collected in the A Sand County Almanac (which should be required reading for any naturalist), Leopold tells of how bright minds are often told to ignore the “music” of nature;

There are men charged with the duty of examining the construction of the plants, animals, and soils which are the instruments of the great orchestra. These men are called professors. Each selects one instrument and spends his life taking it apart and describing its strings and sounding boards. This process of dismemberment is called research. The place for the dismemberment is called a university.

A professor may pluck the strings of this own instrument, but never that of another, and if he listens for music he must never admit it to his fellows or to his students. For all are restrained by an ironbound taboo which decrees that the construction of instruments is the domain of science, while the detection of harmony is the domain of poets.

Professors serve science and science serves progress. It serves progress so well that many of the more intricate instruments are stepped upon and broken in the rush to spread progress to all backward lands. One by one the parts are thus stricken from the songs of songs. If the professor is able to classify each instrument before it is broken, he is well content.

Science contributes moral as well as material blessings to the world. The great moral contribution is objectivity, or the scientific point of view. This means doubting everything except facts; it means hewing to the facts, let the chips fall where they may. One of the facts hewn to by science is that every river needs more people, and all people need more inventions, and hence more science; the good life depends on the indefinite extension of this chain of logic. That the good life on any river may likewise depend on the perception of its music, and the preservation of some music to perceive, is a form of doubt not yet entertained by science.

It would be a mistake to paint all practicing scientists with such a broad brush, but the danger of becoming so objective that the melodies of songbirds and the soft rushing of streams become muted is a very real one. This is strange, especially because it was from wonder that science was born, an attempt to explain what had hitherto been subjugated beneath superstition and religion as it exists, not how we may wish it to be. Still, despite the move away from superstition and natural theology, especially since the writings of Darwin came crashing into the public consciousness, religion attempts to retain a hold on “the birds of the air,” “the beasts of the field,” and “every thing that creepeth upon the earth.” In a recent post on the evolution/creationism debate, the author of the blog Doxoblogy opined;

I’ve got only this to say…looking at creation will inevitably point you to an ‘eternal power and divine nature’ that exists beyond us. Looking in Scripture will introduce this ‘eternal power and divine nature’ to you as the Creator God to whom we owe our love and worship (Romans 1:16-2:11). The God of creation is also the God of Scripture and He has a Son, Jesus.

Clearly there is a sense of awe operating here, and there are seemingly countless flash-animated greeting cards, books, videos, and other resources enforcing the notion that every aggregate of soil, blade of grass, or molecule of water practically screams that life was created by the Judeo-Christian God of the Bible. Such arguments have even become politically fashionable, allowing current presidential candidate John McCain, in an attempt to eat his cake and have it too, to say “I believe in evolution. But I also believe, when I hike the Grand Canyon and see it at sunset, that the hand of God is there also” when asked if he “believed” in evolution. Such a notion clearly points to subjective notions of beauty, as McCain did not say “When I look at a lamprey or a hagfish, I see the hand of God at work.” The overall association of God with the aesthetically pleasing could explain why Thomas Kinkade paintings, which differ so little that I can scarcely tell one from another, as a staple in evangelical Christian households. Perhaps there are some on the fringe that would prefer to think of a tapeworm or liver fluke when contemplating the glory of God, but the vast majority of creeping, crawling, sucking, oozing, and pulsating things on the planet are not generally thought of as being “first in the ways of God.” Even Darwin expressed his doubts about a Creator that was seemingly so cruel. In a famous letter to the American botanist Asa Gray, Darwin confided;

I am bewildered. I had no intention to write atheistically. But I own that I cannot see as plainly as others do, and as I should wish to do, evidence of design and beneficence on all sides of us. There seems to me too much misery in the world. I cannot persuade myself that a beneficent and omnipotent God would have designedly created the Ichneumonidae [parasitic wasps] with the express intention of their [larvae] feeding within the living bodies of Caterpillars, or that a cat should play with mice.

While generally forgotten today, the view that nature is filled with goodness or pleasurable things was, as far as I currently understand, put forth by William Paley in Natural Theology, one of Paley’s core beliefs being that the ability to feel pleasure (and not just pain) was evidence of a beneficent Creator. Still, the understanding that Darwin helped usher humanity, albeit kicking and screaming, into was an understanding of nature that is neither inherently good nor evil. The world is not “for” mankind just as it is not expressly for the benefit of water beetles, Cape Buffalo, or the Northern Flicker. If it were otherwise the world could perhaps divided into creatures that were “good” and others that were “evil,” but no clear distinction exists nor has it ever; attributing such labels to the world around us only speaks to our own ignorance and hubris.

Sometimes I have to wonder if those who propose to have seen God in nature have truly spent any time out in nature or studying its diversity. A poster of some far-flung locale at sunset with one of the Psalm’s printed on the bottom is not understanding nature, a motley organization of life that (try as we might) we are still very much a part of, and even the most repulsive or disgusting of creatures has a worth that does not rely on our “refined” tastes. When an animal dies, insects and bacteria take advantage of the bonanza, putrefying and decomposing the body , beginning the process that will return the creature to the earth. If special circumstances occur, it may well see the light of day again as a fossil, but more likely than not it was be completely broken up, the accumulation of energy and elements in its body being transferred into other organisms and into the ground, allowing different forms of life to flourish. This does not make a maggoty, decomposing carcass any more attractive (or smell any more fragrant), but if we divorce it from our rather superficial requirements of beauty, we can gain an understanding of nature that previously eluded us.

Perhaps my words are only those of a young man, “green” in terms of experience and landlocked in a land of impervious surface and strip malls. Such inexperience may hinder my perceptions, but when I look closely at nature I see neither angels nor demons, God and the Devil being absent from the crashing of the waves along the shore or the lighting strikes of a late-August thunderstorm. This does not mean, however, that I view nature divorced from any sense of awe or deeper emotional feeling, and I would imagine that many of my readers here would tell you the same. When I was covered in wet muck of the Inversand Marl Pit, my heart skipped a beat as a pulled out a chocolate-colored bone fragment that had been kept snug in the greensand for at least 65 million years; my mind reeled at what I could have discovered (and what else might remain buried), and I had to hold back my excitement as I asked my professor if I had really found bone or not. Any book describing the adventures and work of a more professional and seasoned paleontologist or field scientist will reveal much the same thing; objectivity is necessary for the sake of accuracy, but it often comes after a rush of excitement or amazement at a new observation or discovery.

This post is a bit of a throw-away, however, as Charles Darwin long-ago succinctly summed up the thesis of my long argument;

There is grandeur in this view of life, with its several powers, having originally been breathed into a few forms or into one; and that, whilst this planet has gone cycling on according to the fixed law of gravity, from so simple a beginning endless forms most beautiful and most wonderful have been, and are being, evolved. – Charles Darwin, On the Origin of Species by Natural Selection

A female gorilla and two babies at the Bronx Zoo

Convergence or Parallel Evolution?

Many of the world’s great natural history museums devote at least one hall to creatures that no longer exist today. In the old tradition, in order to keep any young upstarts from getting any ideas about evolution, skeletons or parts of skeletons were grouped by the functions they performed, a visitor being likely to find the wing of a bat and the wing of the bird in the same display case even though the two animals extremely distantly related. Newer layouts, conversely, have largely ignored the end-function of one line or another to group animals together by homology and their shared characters, the most well-known example being the remodeled 4th Floor of the American Museum of Natural History in New York City which has attempted to arrange its fossil collections as a walk-through cladistic diagram.

Still, the generally discarded of grouping animals by their adaptations to general habitats or niches is not without it’s charms. Over and over again, evolution has produced forms that seem to converge on certain body plans, varying habitats making some traits advantageous and others a liability, helping to adapt different organisms to their local ecologies. Flight has independently evolved several times (and the ability to glide an even greater number of times), as well as adaptations to marine environments, saber-like canine teeth, immense sails along the spine, and slicing premolar teeth, although each time such familiar features seem to arise it shows that there is more than one way to solve an evolutionary problem from any given point in an organism’s natural history. Not everything can be chalked up to convergence of form in order to carry out particular functions, however. Parallel evolution, although sometimes difficult to determine, also allows relatively closely related forms to take the same evolutionary paths, showing many of the same anatomical characters even though they diverged from a common ancestor at some point in the past and occupy at least two different lines of descent. In fact, it is often these weird and wonderful creatures that are forgotten or overlooked, more people recognizing the term “saber-toothed cat” (or, loathe as I am to say it, “saber-toothed tiger”) or the genus Smilodon than the term “Nimravid” or the genus Dinictis. The following entry, therefore, will be an attempt to navigate through the somewhat “entangled bank” of evolutionary relationships among animals that appear to be shaped in similar ways by the environment but constrained by their species’ history, showing us that there is more than one way to make a saber-toothed cat.

Back into the pool: Of Ichthyosaurs, Sharks, and Cetaceans

Perhaps one of the most well-known (or at least widely cited) examples of evolutionary convergence has been that of the similar body shapes of sharks, ichthyosaurs, and cetaceans. It’s difficult to see these three distinct groups of creatures side by side and not recognize the similarities, but why are they similar in the first place? If they belong to groups that are distantly-related branches of the evolutionary “bush,” why should they have developed similar body forms?

One of the most well-known examples of evolutionary convergence; (From Top to Bottom) An ichthyosaur Ophthalmosaurus icenicus, a Porpoise, and a Spiny Dogfish (Squalus acanthias)

From the 1925 creationist book The Predicament of Evolution by George McReady Price.

Creationists have been quick to seize upon the idea of convergence as if it were one of evolution’s weak points. In 1926, George McCready Price wrote the following in one of the more well-known early American anti-evolution texts, The Predicament of Evolution;

For instance, we have the shark, the ichthyosaur (an extinct kind of fish-shaped reptile), and the dolphin (a true warmblooded mammal, and not a fish at all), all of which greatly resemble each other in external shape and general appearance. Each has the same long, sharp snout, the same powerful tail, the same general fishlike shape. And yet the first of these is a true fish, the second was just as true a reptile, while the third is a mam-mal, bringing forth its young alive and feeding them by milk, just as does a cow or a horse, though it lives in the sea.

Here the evolutionists have to say that this peculiar shape and general form has been evolved separately and independently in each of these three instances. Indeed, Henry Fairfield Osborn, President of the American Museum of Natural History, New York City, declares that a very similar shape and form has been independently evolved “at least twenty-four times.”—”Encyc. Brit.,” Vol. XX, p. 578…

From this large group of facts we become convinced that these many similar or identical structures, which must have been evolved quite independently (if evolved at all), make too great a draft on our credulity. At least, these hundreds of examples of “parallel evolution” greatly weaken our confidence in homology, or similarity of parts and organs, as a proof of blood relationship.

Such arguments have become traditional amongst creationist apologists, suggesting that if convergent evolution does occur then we must throw homology out the window as similar structures will only mislead us as to the true affinities of the creatures being studied. As we will later see with Cuvier’s Ptero-dactyl, this can be a danger for scientists who are unwary and wish to shoehorn creatures into existing taxonomic categories, but not for those who actually look beyond superficial appearances.

The reason why the shark, the ichthyosaur, and the porpoise should all look vaguely the same is because they live(d) in the same environment; the ocean. An organism that is suspended in a fluid that is much denser than air can be adapted in various ways to such an “alien” environment, but physics does dictate what shapes can be taken based upon life history. It is possible to be a floating filter feeder, exhibiting a round shape, but such a strategy is essentially out of the question for animals that need to move quickly and to hunt for food. What is required is not only a powerful propulsive organ to keep the organism moving forward, but also extra appendages to allow for the control of movement and a streamlined shape to reduce drag (and hence reduce energy costs for moving through the water).

One of Charles R. Knight’s renditions of an ichthyosaur.

In fact, sharks as a whole provide a good model for various forms of ichthyosaurs. While ichthyosaurs are generally presented as already being streamlined and possessing a large caudal fin with two equally long lobes, we would be loathe to forget that they too are products of evolution and many fossils show us that they were not always an Euryapsid (thank you, johannes) answer to modern-day Lamnid sharks. Early ichtyhosaurs actually had more of a “bump” towards the back of their tail rather than a full-blown caudal fin, their overall body shape and lack of a large propulsive surface keeping them from moving too quickly through the water. A similar tail type/form can be seen in many modern day sharks like the Nurse Shark, which generally live along the bottom feeding on crustaceans and inhabitants that can be sucked out of coral crevices. Being that ichthyosaurs lack gills, it is unlikely that their early representatives were bottom-dwellers, instead preferring shallow areas, which can be especially productive in terms of food.

Modification of the “tail kink” (which was at first thought to be a taphonomic feature, early reconstructions showing “amphibious” ichthyosaurs with straight tails) seen in early forms allowed for the eventual evolution of a crescent-moon shaped tail, as well as adaptations in the skull and of the limbs into fins (the addition of digits and the addition of bones in the digits being quite common in the latest forms). This more-familiar shape would allow ichthyosaurs maximum propulsion with their caudal fin (the spine going downwards instead of upwards, as in sharks) while they would be able to exert control over their motions with their pectoral fins and would be kept from rolling in the water by their dorsal fins. The evolution of large eyes and other features aside, the overall shape and basic skeletal structure of ichthyosaurs seems to be an optimal design for medium-to-large, fast-moving, oceanic predators (although mosasaurs, pliosaurs, and plesiosaurs took different evolutionary routes).

What allowed ichthyosaurs to develop an effective side-to-side motion of the tail would not work for cetaceans, however. Ichthyosaurs developed their mode of propulsion by side-to-side motions of the spine, perhaps swimming in a mode similar to eels or cat sharks at first, a common form of locomotion in modern reptiles. This sort of motion is usually accomplished on land via a sprawling gait, the limbs being held out to the sides and the animal exhibiting a bit of a side-to-side motion as it moves along.

Whether the ancestors of icthyosaurs were sprawlers (to a greater or lesser extent, predisposing them to side-to-side motions of the tail and body) or not, cetaceans evolved much more recently in evolutionary history, and developed from ancestors that carried their legs directly underneath their body. The plasticity of early archaeocetes and their artiodactyl ancestors was greatly diminished, their hip and spine structure adapted to up-and-down undulations rather than the side-to-side motion seen in the video of the salamander. This sort of constraint has not stopped mammals from becoming adapted to the water, however, and clues to the evolution of cetacean movement can be seen in living animals like Giant River Otters;

In the water, undulations of the spine accompanied with some propulsion from the limbs proves to be very effective, and it’s not hard to imagine an archaeocete like Ambulocetus, as my friend Neil so aptly described, as a “sexy otter.” Once undulation of the spine became established as a method of moving through the water, the eventual addition of a tail fluke would do for cetaceans what the crescent-shaped tail of tuna, sharks, and icthyosaurs acheived in terms of speed and power, the body being adapted towards a streamlined appearance with (again) the pectoral fins providing lift/control and the dorsal fin preventing rolling. Larger forms of whales, namely the Mysticetes or Baleen Whales, grew to immense size and gave up some of the features that seem to be convergent with sharks and the smaller ichthyosaurs (some, in fact, did acheive whale-size), but they are derived from more predatory designs and their niche as massive, far-ranging suspension feeders free them from some constrains while imposing some new ones.

A painting of leaping ichthyosaurs by Heinrich Harder (circa 1916)

Human engineering has recognized similar constraints for motion in the water and even in the air; planes and submarines most closely resemble sharks and dolphins in overall shape, the placement and size of the wings on a 747 having much the same function as the large pectoral fins of far-ranging pelagic fish like the Blue Shark. Life in the water adapted all three groups of animals towards the same shape because there does not seem to be any other way to be a fast-moving, medium-to-large sized marine predator; speed and some degree of maneuverability are paramount. Some other lines have diverged from this shape, as noted before, but the sharks, dolphins, and (I don’t think it’s too much of a stretch to say) ichthyosaurs all occupied essentially the same niche and therefore were adapted in a particular fashion.

Do not think, however, that the convergence of three lines towards one body plan gives credence to a kind of “orthogenesis” or progressive force driving evolution. There was no sort of supernatural or external force manipulating the genetic material of these groups with the shape of a dolphin or shark in mind. Rather, the environment and local ecology determined what form would be favored through time, and even though the three groups may look the same and have significant convergences, they also have many traits in common with their ancestors, allowing us to trace their evolutionary history (which is why no one is arguing that dolphins, sharks, and ichthyosaurs are closely related or form a small monophyletic grouping).

A marsupial you wouldn’t want to meet

Living members of the Carnivora (bears, cats, dogs, civets, weasels, etc.) have always caught my attention, but there was an entire group of carnivorous mammals, now extinct, that have left no living representatives. The last known member of this group was named Thylacoleo carnifex by Richard Owen, and it has some of the strangest dentition ever seen in a marsupial. Marsupial mammals are well-known in Australia, creatures like kangaroos, koalas, and wombats coming most immediately to mind out of living extant taxa. There was a much more diverse population of marsupials during the Pleistocene, however, and the “marsupial lion” was likely a formidable predator.

A skull of Thylacoleo on display at the AMNH.

In order to understand why Thylacoleo is relevant to our discussion of convergence we need to first understand what makes living placental Carnivores so special. Many carnivores, especially cats, have a rather specialized dentition, certain molars and premolars making up what is known as the “carnissal shear.” These teeth are pointed and act like scissors, easily cutting up flesh or crushing bone. The molars behind the shear are often reduced (some groups have retained their molars in order to incorporate a more generalized diet, like dogs and bears), the dental specialization perhaps being one of the keys to the success of this group. Earlier predators of now-extinct lines like Mesonychids lacked such specialized cutting teeth, and the teeth behind the canines of the large Andrewsarchus show that their oral tool-kit was a bit more blunted.

The skull of Andrewsarchus, on display at the AMNH

Thylacoleo, a carnivorous marsupial not descended from the Miacids that gave rise to living carnivores, also developed something of a “carnissal shear” but in a different way. Rather than a battery of teeth that became sharpened, one of the upper and lower premolars of Thylacoleo became elongated and blade-like, and the cleaver-like teeth helped to sharpen each other as they moved past each other when opening or closing the jaw. Thylacoleo also had a terrible bite, the attachments for the muscles that opened and shut the jaw were massive, somewhat constricting the amount of space the brain could take up, but giving Thylacoleo what was perhaps the most powerful bite forces amongst mammalian predators, especially given it’s relatively small body size (it was only about four feet long and 220 pounds).

Thylacoleo is an odd marsupial in another respect; the claw on its thumb was retractable like that of a big cat. This sort of adaptation is especially useful in keeping claws sharp, and perhaps keeping the claws sharp would allow Thylacoleo to get a good hold on its prey before going to work on it with its teeth. At this point I should probably mention that some scholars in the past have thought that Thylacoleo was an herbivore, not unlike the extant marsupial Phalangers. I will leave the response to such an argument to Richard Owen;

These eminent authors received the support, in reference to objections to my conclusions, of the (then) Curator of the Australian Museum, Sydney, Mr. GERARD KREFFT, who, in his contribution to the ‘Annals and Magazine of Natural History,’ series 3, vol. 18, 1866, p. 148, records his opinion that “the famous marsupial Lion was not much more carnivorous than the Phalangers of the present time.”

The species of carnivorous Phalanger is not named. No evidence of such by fossil specimens has reached me, nor have I found such exceptional habit of an existing species of Phalangista elsewhere noted.

As my friend Zach has noted, however, calling Thylacoleo a “marsupial lion” is a bit misleading. Even though some lion-like aspects of the skull (the results of convergence on a hypercarnivorous lifestyle, and Thylacoleo means “pouched lion”) led the anatomist Richard Owen to name the creature on the basis of such resemblances, the ways in which Thylacoleo shows its marsupial affinities are much more important. Referring to this animal as the “marsupial lion” without qualifications (as well as calling the extinct Tasmanian Tiger the “marsupial wolf”) usually confuses more than illuminates, and creationists often take the names and superficial resemblances to mean that evolution didn’t occur. Instead, they propose that God made the beginning of a “kind” of carnivorous mammal which was preserved on Noah’s Ark and gave rise to all later forms, important reproductive habits deemed to be of little consequence.

Even so, Thylacoleo carnifex and its relatives represent a branch of marsupials that became fairly specialized predators, and given the plasticity of tooth structure, it’s not hard to see how sharp premolars could be adapted into a blade to cut flesh. While it may be easy to draw connections between this animal and living carnivores, however, perhaps we should be more measured in our descriptions; both groups met the same challenges in similar ways, but the differences are far more striking and important in this example of convergence on a particular niche.

On what day were the Ptero-Bats created?

An engraving of the creature now known as Pterodactylus antiquus, the very one described by Collini.

Before there were natural history museums, there were motley assortments of organic odds and ends known as curiosity cabinets, and in the cabinet of Karl Theodor there would eventually come to be a petrified treasure. Although it was probably collected around 1767, the first known pterosaur fossil was not described until 1784, when the appointed caretaker of the collection, Cosimo Alessandro Collini, attempted to determine the nature of the strange creature that came to him from the limestone of Bavaria (the same deposits that later yeilded Archaeopteryx). Although certain that he was the remains of an animal from an earlier time, Collini was agnostic about what kind of animal he had come to possess. Years later, the famed anatomist Georges Cuvier investigated Collini’s paper and illustrations, noting that the creature was certainly a reptile. Still, the fossil would remain without a proper name until Cuvier would write a more detailed analysis in 1809, dubbing the fossil “Ptero-dactyle.”

Not everyone agreed with the analysis of Cuvier, however, especially since Cuvier did not get to see the fossil himself and had to work from the drawings in Collini’s paper. Samuel Thomas von Soemmerring, of the Bavarian Academy of Science, thought that the pterosaur was some unknown type of bat, a view that would remain entrenched in the minds of some scientists for many years. Indeed, one restoration by Edward Newmann in 1843 (and “re-drawn” for Gosse’s work Omphalos, as shown below), depicted the two known types of pterodactyl known at that time as fuzzy bats, complete with cute little ears. It is clear from the drawing that pterosaurs do not make good bats, although this didn’t stop many German paleontologists from taking such a stance through the first half of the 19th century.

Newmann’s “marsupial bats”, conspicuously missing their ears, from Gosse’s Omphalos. It’s likely that Gosse recognized the reptilian nature of these Pterodactyl by the time he wrote his book, so Newmann’s work was copied minus the more mammalian aspects.

But why was there such confusion? It is likely because there is something familiar about pterosaurs that had been seen in living bats; the extension of digits to hold a membraneous wing. While the first fossil, despite wonderful preservation, did not preserve a membrane impression, it is hard to look at it and not recognize the superficially similar structure of a bat’s wing, which also carries a membrane to enable flight. In fact, birds seem a bit unusual in developing feathers for flight; many varieties of gliding and flying creatures have taken to the air (regardless of whether they engage in powered flight or glide) by the use of membranes. Indeed, gliding may often precede powered flight, and once an animal has developed a membrane that can be stretched between its limbs to glide, the extension of the digits at the point(s) of attachment can help to expand the wing size. Such changes likely occur as a result of changes in development, natural selection favoring the invasion of a new niche based upon variations that exist in a population, although in the case of pterosaurs we can no longer test to see if this is correct.

As we just saw with Thylacoleo, however, the convergences of pterosaurs and bats are rather slight, overall. While both acheived flight on membraneous wings attached to extended digits (many more in the case of bats) and have relatively compressed bodies, pterosaurs had a much greater diversity in shape and size than modern bats. Likewise, they did not elongate the rest of their fingers, suggesting that there was some situation (be it climbing or hanging on to a perch) that the pterosaurs still needed their other fingers for (although bats can climb pretty well with their thumbs, and some have even evolved suction disks). Still, it can be said that both took to the air by similar means and had to deal with similar constraints, but their evolutionary paths are far more divergent than that of the aforementioned sharks, ichthyosaurs, and cetaceans.

It doesn’t look like much of a planet-eater to me

A female Gharial at the National Zoo in Washington, D.C.

Perhaps one of the most unnecessarily confusing groups of extinct animals are the phytosaurs. Filling the niche now occupied by reptiles like the Saltwater Crocodile, the water-dwelling archosaurs have left no living descendants despite their past diversity. At first glance, the phytosaur Rutidon looks just like a modern-day Gharial, and even though it shares a common ancestor with the reptiles that now exist in tropical watery habitats all over the world, it is not otherwise related. The most prominent phytosaur feature is that their noses are over or just anterior to their eyes on their head, not at the end of their snouts. This would allow them to breathe while completely submerged, although their eyes might not have been above water when hiding in such a manner. Even beyond this feature, their jaws seem to be fairly simple, merely having a hinge at the back to open-and-close. Compare this arrangement, here represented by the giant Machaeroprosopus gregorii, with the more complex reconstruction of the true crocodilian Deinosuchus (although, admittedly, this reconstruction was heavily based upon the living Cuban Crocodile and may not be fully accurate. It still serves to show the differences between the groups, however).

Machaeroprosopus, currently on display at the AMNH

Deinosuchus reconstruction, formerly on display at the AMNH

The most notable difference are the complex bones at the back of the throat of Deinosuchus which are arranged to slide past each other as the jaw opened and closed. No such feature is seen in the giant phytosaur. Still, even after the phytosaurs died out, crocodilians did not return to the water until about the Cretaceous period, many forms being absolutely terrifying land predators that have also long been extinct. One of the early forms was Protosuchus, a small true crocodilian that represented a line that changed little during its tenure on the earth.

Reconstruction of Protosuchus

Outside of walking relatively high off the ground, Protosuchus had a foreshortened snout which was lower than its eyes, quite different from the arrangement in living crocodilians. As seen in the Dwarf Caiman photograph, below, living crocodilians have their eye sockets on the top of their head, their eyes sticking out on the surface as well as the tip of their nose when they lie in wait for prey (or just rest, for those who would like a less sensationalist tone). Protosuchus, by contrast, has eyes to the sides of the head, even facing somewhat forward, showing that it was much more well-adapted to the land than any swamp or shallow pool. Crocodilans did eventually enter the water, however, and their fossils are among the most common of any vertebrates. Some, like New Jersey’s very own Thoracosaurus, even became marine species, and a few varieties evolved crescent-shaped caudal fins on the ends of their tails to help them swim. The common belief, however, is that crocodiles have always been crocodiles, “changing little since the time of the dinosaurs,” and such generalized half-truths do little justice to crocodilians or their distant phytosaur cousins.

Saber-toothed Nimravid doesn’t sound quite the same…

Many museums have cases devoted to the great saber-toothed cats of epochs long gone, but it would take someone with more than just a cursory understanding of paleontology to sort out what is really being displayed. Saber-teeth, or elongated canines, have evolved many times over in the course of mammalian history, showing up in herbivores like the living Musk Deer as well as extinct groups like the gorgonopsids. Animals as different as a Musk Deer and Inostrancevia are fairly easy to tell apart, even for the non-specialist, but what about nimravids and the “true” saber-toothed cats?

A diagram of the three ideas of Nimravid/Felid evolution.

James Whitcomb Riley is purported to have once written “When I see a bird that walks like a duck and swims like a duck and quacks like a duck, I call that bird a duck.” Unfortunately, this argument is quite popular (even being utilized by the likes of prominent Intelligent Design advocate Michael Behe) despite being very superficial and even vapid. Needless to say, it doesn’t apply to our discussion of Nimravids and true felid saber-toothed cats, but in decades past the two groups were lumped together.

<img src=”” alt=”Skulls” />

So, what makes a nimravid a nimravid? They look awfully like cats, so why aren’t they included in the Family Felidae? What makes such distinctions so difficult is that those investigating the skull of Smilodon and Eusmilus would have to be relatively well-versed in scientific jargon and anatomy in order to point out the most important differences. While some nimravids (like Eusmilus) had large canines, their teeth alone are not diagnostic, and the original factors used by E.D. Cope that differentiated these animals from “true” cats were the “alisphenoid canal, postglenoid foramen, carotid, posterior lacerate, and condyloid foramina, postparietal foramina” in the skull (Hunt, 1987). The various canals and foramina listed dictate the paths of various nerves and blood vessels in the skull, and the arrangement in nimravid skulls seem to be more primitive compared with true felids. Likewise, nimravids lack a two-chambered auditory bulla, which is a rounded bit of bone associated with the ear which true cats posess.

There are a few more obvious giveaways when dealing with some nimravids, however. Nimravids equipped with long canines often have more cone-shaped canines than those of saber-toothed cats (which are flatter in cross-section), and many have bony “sheaths” extending from the lower jaw into which the massive teeth fit. Perhaps the most famous example of this kind of arrangement is the genus Barbourofelis, an animal that has actually been assigned to its own family as it is likely more closely related to true cats than nimravids (Barbourofelis was previously classified as a nimravid). Because of this (and the fact that another cat-like offshoot, the marsupial Thylacosmilus) the tooth-sheath shouldn’t be considered diagnostic of nimravids only, but it does give you a substantial clue that you’re probably not dealing with an actual saber-toothed felid.

Despite these differences, it has often been difficult to differentiate the groups (and debate still continues). The diagram above, based upon one in Robert Hunt’s 1987 paper “Evolution of the aeluroid Carnivora. Significance of auditory structure in the nimravid cat Dinictis,” offers three simplified versions of the hypotheses about the relationships of nimravids and felids. Initially it was thought that there was a progressive evolution from ancestor to descendant in a straight line, the nimravids being the direct ancestors to the saber-toothed cats. This view does not represent how evolution truly works, however, and was found to be incorrect. In its place came a view that nimravids and saber-toothed cats diverged from a common ancestor at about the same time, going off in separate directions. This is better, and is more consonant with the data, but again it suggests that the line representing the common ancestor went extinct, either in becoming nimravids or saber-toothed cats. What seems to be the case based upon current data is that the nimravids split off from a common ancestor somewhat before the saber-toothed cats, the line containing their common ancestor continuing its own evolution as both groups evolved. Such a branching pattern is not unusual, and should even be expected, especially since there are living primates like tarsiers and lemurs that represent the overall kind of animal our ancestors once were, but still quite different and undergoing their own evolution alongside our own lineage.

The skull of Thylacosmilus, the marsupial answer to the saber-toothed cat, on display at the American Museum of Natural History in New York City. Note how far back in the skull the roots of its massive canines extend.

The skull of Megantereon on display at the AMNH. It was one of the “true” saber-toothed cats.

To complicate things even further, the skull or skeleton of the marsupial Thylacosmilus is also often thrown into the mix. Although totally unrelated to nimravids or felid saber-toothed cats, the South-American Thylacosmilus converges closely on the appearance of the placental predators, although there are some important differences. As can be seen from the above photographs, the eye of Thylacosmilus is entirely enclosed by bone on the side of the head, while in many felids and nimravids the eye socket is not entirely ringed-in by bone as if someone had bored a hole in the skull (compare the skull of Thylacosmilus with that of Thylacoleo, above). Further, the teeth of Thylacosmilus have very deep roots, going back in the skull almost over the eye. Originally it was thought that the teeth faced outward, but this was based upon a distorted skull and later finds showed the true position of the long canines.

Hoplophoneus

Now that we have elimated Thylacosmilus from the running as another case of marsupial convergence, we must ask why the nimravids and felid saber-toothed cats are so close to each other in appearance. While many of the instances I’ve discussed previously have been instances of convergence, be it throughout the entire body or merely certain aspects of it, the nimravid-felid connection is a wonerful example of parallel evolution. W.E. le Gros Clark provides an excellent summation of understanding the difference in his 1959 book The Antecedents of Man;

From what has already been said, it is clear that, in assessing degrees of phylogenetic affinity, it is always necessary to take into account the factors of parallelism and convergence in the evolutionary development of related or unrelated groups. These processes can lead to structural similarities, which, taken by themselves, may be misleading. The term convergence is applied to the occasion in general proportions or in the development of analogous adaptations in response to similar functional needs. But such similarities are superficial and easily distinguishable by a detailed comparative study of the animal as a whole. For example, the resemblance in general appearance, even in a number of morphological features, of the Tasmanian wolf to a dog does not obscure the fact that in fundamental details of their anatomical construction they belong to quite different mammalian groups. On the other hand, the potentialities of parallelism seem often to have been much overestimated by some anatomists, for this phenomenon has sometimes been invoked in support of extreme claims for independant evolution of groups which are almost certainly quite closely related. We can agree with G.G. Simpson that the whole basis of parallelism depends on an initial similarity of structure and the inheritance of a common potentiality for reproducing homologous mutations, and that, this being so, the initial similarity and the homology of mutations themselves imply an evolutionary relationship. Expressed in another way, it may be said that convergence increases resemblances (which are, however, no more than superficial), while parallelism does not so much increase resemblances as maintain and perpetuate (by development ‘in parallel’ so to speak) similarities which have already existed ab initio in the genetic make-up of related types. Thus, ‘closeness of parallelism tends to be proportional to closeness of affinity.’

There are a few problems with this reasoning, namely that it seems to give credence to an almost pre-determined genetic course for the lines to evolve in parallel, although le Gros Clark makes it clear in the work that he does not support in any way the notion of orthogenesis. Still, the passage makes the important distinction that in order to undergo parallel evolution groups need to be somewhat closely related and already bear similar structures, evolution preserving many of the similar traits instead of working to the same end from two disparate points. In the case of the nimravids and the felids it seems that they evolved from a common ancestor which was probably taken to carnivory. Nimravids branched off earlier, being more “primitive,” while the felids came off the same line (or a very similar one) after it had accumulated a few more evolutionary changes. Indeed, even if form seems to be static or change little, it’s hard for me to believe that designs are not slightly adapted this way or that as if the creature was an already perfect creation not influenced by changing ecological circumstances. Still, it seems that the nimravids and felids were adapted in similar ways, their ancestral lines probably possessing at least semi-retractable claws, long and sharp canines (although not long to the extreme like its descendants), a shortened face, and a developed carnissal shear. It is really not that difficult to change a civet-like creature (or in the case of our hypothetical common ancestor, a creature a bit closer to a cat) into a saber-toothed Smilodon, the changes being modifications of existing structures more than the creation of something entirely new out of nowhere. In fact, the vertebrate tetrapod skeleton has proven to be quite versatile, and most of the major bones in any vertebrate skeleton can be found to correspond with those in another vertebrate, allowing us to compare rhinos with ceratopsians, dromeosaurs with birds, cats with dogs, ichthyosaurs with cetaceans, and humans with primates.

Of constraints and convergence

I hope that is has become clear why convergence is such a strong theme in the evolution of vertebrates. At this point in the history of evolution, vertebrates have had a chance to fill nearly every niche imaginable in a large variety of habitats over millions of years, and so common themes are bound to arise. When groups return to the ocean, the environmental constraints shape them in ways peculiar to their new way of life that would not be advantageous in other situtations (i.e. being such a large aquatic animal that you’d be crushed by your own weight if you came onto land). When mammals become adapted to be predators, their dentition and morphology must be altered if they are to be successful hunters, carnivores past and present showing some suprising similarities despite being only distantly related. Even when taking to the air, laws of physics still apply, and natural selection often works through physical and chemical constraints to produce new forms.

It is of little doubt that the tetrapod design is a versatile one, retaining its overall character through the various changes that it has endured. Indeed, even when a lineage dies out and may seem gone forever, there is no law that says a similar situtation in the future will not produce forms that may be strikingly familiar, even if such organisms are not directly related to the last group that filled their new niche. Evolution has produced “endless forms most beautiful and most wonderful” and will continue to do so long after I am gone, but random mutation/natural selection do not work in isolation from the rest of the natural world. Evolution has produced so many amazing creatures precisely because ecology, physics, and chemistry have offered up both opportunities and challenges, and I only regret that I will not be able to witness the familiar and unfamiliar about what is swimming in the seas, flying in the air, or stalking the land 500 million years from now.

Scuttling the Aquatic Ape Hypothesis

On our occasional trips to the New Jersey shore, my wife is always the first one in the water. While I’m cautiously wading in, dreading that final slap of cold water just below my waist, she’s already frolicking in the waves, egging me on to just jump in and get it over with. Eventually I too become submerged (either willfully or by force of a wave I never saw coming), salt water inevitably shooting up my nose. Don’t get me wrong, I do enjoy warm days at the beach, but on each visit it seems that I as an individual, if not a representative of a population or species, am not well-adapted to a near-shore marine existence. Followers of the Aquatic Ape Hypothesis* (AAH), however, beg to differ.

[* I say “hypothesis” and not “theory” (AAT) because the writings of Elaine Morgan and others do not have enough supporting evidence to garner it the more prestigious title of “theory.” Given the current paucity of evidence and research, the Aquatic Ape Hypothesis is precisely that and no more.]

Before discussing the current manifestation of the AAH, we need to go back to a time when the truth of evolution had yet to fully take hold in the minds of scientists and philosophers. The Ionian philosopher Anaximander (610-546 BCE), student of Thales, suggested that the world first existed in an entirely aquatic state, the recession of the globe-consuming waters creating life. In From the Greeks to Darwin (1905), famed American Museum of Natural History president Henry Fairfield Osborn described the views of Anaximander as follows (a similar treatment is given in Osborn’s Man Rises to Parnassus, as well);

He conceived of the earth as first existing in a fluid state. From its gradual drying up all living creatures were produced, beginning with men. These aquatic men first appeared in the form of fishes in the water, and they emerged from this element only after they had progressed so far as to be able to further develop and sustain themselves upon land. This is rather analogous to the bursting of a chrysalis, then to progressive development from a simpler to a more advanced structure by a change of organs, yet a germ of the Evolution idea is found here.

We find that Anaximander advanced some reasons for this view. He pointed to man’s long helplessness after birth as one of the proofs that he cannot be in his original condition. His hypothetical ancestors of man were supposed to be first encased in horny capsules, floating and feeding in water; as soon as these ‘fish-men’ were in a condition to emerge, they came on land, the capsule burst, and they took their human form.

Like the works of many Ionian philosophers, the ideas and opinions of Anaximander do not seem to have taken hold (Aristotle ultimately becoming the preferred scientific and philosophical source for further consideration in Europe in centuries to come), and not much of his work remains. It is curious to note, though, that wrong as Anaximander was about the origins of humans, the reasons he uses to support his ideas (as relayed by Osborn) are very similar in approach to those of Elaine Morgan and some modern-day AAH adherents, as we shall soon see.

To the best of my current understanding, the hypothesis that man was a product of the sea did surface again until 1942 when Max Westenhofer of the University of Berlin published the book The Unique Road to Man. According to Donna Kossy’s book Strange Creations, Westenhofer’s treatment of an aquatic origin of mankind consisted of little more than mention of it as a promising hypothesis, however, and the outbreak of war prevented the professor from pursuing the line of inquiry further. The hypothesis would have to wait until March 5, 1960, when marine biologist Sir Alister Hardy presented a lecture on “Aquatic Man: Past, Present, and Future” to the British Sub-Aqua Club. The address caused quite a stir and led Hardy, who had been inspired by the layers of sub-cutaneous present in humans and some marine mammals he had seen skinned on a journey to the Antarctic in 1927, to write a series of articles in the magazine New Scientist to clarify his position on the subject. Kossy relates the words of Hardy from an April issue of the magazine (although the year is not specified);

My thesis is that a branch of this primitive ape-stock was forced by competition from life in the trees to feed on the seashores and to hunt for food, shell fish, sea-urchins, etc., in the shallow waters of the coast. I suppose that they were forced into the water just as we have seen happen in so many other groups of terrestrial animals. I am imagining this happening in the warmer parts of the world, in the tropical seas where Man could stand being in the water for relatively long periods, that is, several hours at a stretch. I imagine him wading, at first perhaps still crouching, almost on all fours, groping about in the water, digging for shellfish, but becoming gradually more adept at swimming. Then, in time, I see him becoming more and more of an aquatic animal going further out from the shore; I see him diving for shell fish, prising out worms, burrowing crabs and bivalves from the sands at the bottom of shallow seas, and breaking open sea-urchins, and then, with increasing skill, capturing fish with his hands.

Thus the more familiar image of the amorphous “Aquatic Ape” was born, wading out into the surf and feeling in the shallow sands for food. The early stage of such a transformation is awfully raccoon-like, as raccoons have incredibly sensitive hands that they use to feel about in streams and shallow waters for mussels, crayfish, and other morsels without being driven to become fully aquatic themselves. Nevertheless, the idea that man had his origins in a shallow sea rather than on a hot and brutal savanna was certainly controversial. Ever since Raymond Dart described the skull of the Taung Child in 1925 (shifting attention away from Europe and Asia for the origins of man) and the fossil assemblages of the South African caves were discovered, humans were thought to have evolved through a hunting culture, nearly every specialization that separates us from living primate relatives being due to our meat-craving societies. Indeed, the remains of Australopithecus found in South African caves (especially the jaw of a 12-year old child whose jaw appeared to have been fractured by a direct and accurate blow) like those Makapansgat suggested to Dart that these “proto-men” were not only skilled hunters, but also murderers and cannibals. Even though our understanding of these assemblages has greatly changed since Dart’s time (see C.K. Brain’s The Hunters or the Hunted?), the overall image of human evolution being intricately linked to meat-eating and hunting has dominated the discussion of our origins. Even more specifically, the considerations of our ancestors have nearly always focused on the male of the species, and even Hardy’s early ideas of an aquatic ape focused primarily on males.

In 1964, zoologist Desmond Morris published the bestseller The Naked Ape. Today the book is nearly useless outside of understanding the history of thought about human evolution, but when it was first published a short discussion of the AAH caught the attention of a woman named Elaine Morgan. On page 37 of the 1967 paperback edition, Morris states;

Another, more ingenious theory is that, before he became a hunting ape, the original ground ape that had left the forests went through a long phase as an aquatic ape. He is envisaged as moving to the tropical sea-shores in search of food. There he will have found shellfish and other sea-shore creatures in comparative abundance, a food supply much richer and more attractive than that on the open plains. At first he will have groped around in the rock pools and the shallow water, but gradually he will have started to swim out to greater depths and dive for food. During this process, it is argued, he will have lost his hair like other mammals that have returned to the sea. Only his head, protruding from the surface of the water, would retain the hairy coat to protect him from the direct glare of the sun. Then, later on, when his tools (originally developed for cracking open shells) became sufficiently advanced, he will have spread away from the cradle of the sea-shore and out into the open land spaces as an emerging hunter.

…

Unfortunately, [searching for fossils in marine or fluvial deposits or further research into the AAH] has yet to be done and, despite its most appealing indirect evidence, the aquatic theory lacks solid support. It neatly accounts for a number of special features, but it demands in exchange the acceptance of a hypothetical major evolutionary phase for which there is no direct evidence. (Even if eventually it does turn out to be true, it will not clash seriously with the general picture of the hunting ape’s evolution out of a ground ape. It will simply mean that the ground ape went through a rather salutary christening ceremony.)

[Emphasis mine]

Elaine Morgan read the brief treatment and qualifications (some of which has been omitted here for the sake of brevity) and wanted to know more about the possibility of our ancestors going through an aquatic stage of evolution. No information seemed to be available, and so Morgan wrote to Hardy in 1970, and he encouraged Morgan to push ahead with her research and desire to write a book about the AAH. The result was the bestselling The Descent of Woman, published in 1972. My copy is a little bit newer than that, being the Bantam 1973 edition, and featuring what appears to be a nude mother and child on the cover. Closer inspection reveals that something isn’t quite right, however; the mother I previously assumed was a representative of Homo sapiens looks like she’s been hit in the face with a frying pan. I didn’t know it at the time, but the text would reveal that the plump, nude, and long haired female on the cover was not drawn from life, but rather was Morgan’s idea of the Australopithecus specimen “Lucy” as Aphrodite.

Morgan’s first book is certainly a unique one, weaving in between “Just-so story” type paleo-fiction and long arguments about the female orgasm, including it’s fallacious mythical status. Indeed, the AAH only seems to occupy the first 1/3 of the book, only cropping up here and there in the following chapters, and receiving only a brief mention in the conclusion. Still, the way Morgan structures her argument in her first book will tell us much about her later works and the rise of the AAH as a popular idea. Early on in the work, we are introduced to a hypothetical female ape, not unlike Proconsul, living in Africa sometime during the Pliocene (~5.3-1.8 million years ago). During this time a hard life trying to find food and avoid predators was becoming even harder, Morgan hypothesizing that a terrible heat wave would change the way of life for many populations of these unnamed apes (from what I understand, however, the climate of the Pliocene began to approach our own and became cooler, drier, and had more seasonal distinctions rather than being a global hot-house).

Morgan’s ape was in a bit of a jam, that’s for sure. The water holes are said to be stalked by hungry cats and food was becoming scarce, and the imaginary female was not as fearsome or powerful as the males in her group. Eventually she was chased into the water by a large cat, and decides that, despite her distaste for water, “the seaside not at all a bad place to be. She found to her delight that almost everything on the beach and in the water was either smaller or more timid than she was herself.” Indeed, Morgan’s ape appeared to have found paradise. While other animals cooked during the “dog days of the Pliocene”, her ape (and by extension the population of apes) found what seems to be a sheltered and peaceful lagoon devoid of predators, scavengers, or other threats from either land or sea. “Leopards don’t come so far into the sea, nor sharks so near to the land,” we are told, and while leopards may not be attracted to water, sharks are well-known for their shallow water hunting habits. Crocodiles are not even considered, nor are stingrays, poisonous urchins, jellyfish, disease, infection, or any of the other biological problems that may come with an aquatic existence or change in ecological setting. The new home of the apes sounds better than Club Med, a watery Eden lacking in devious serpents and forbidden fruit.

As suggested by Morris and Hardy, the population of apes gets by on a diet of shellfish and relatively stupid sirenians that happen to come by, males making short work of the water-going creatures with rocks found along the shore. Given the amount of time that the apes would be spending in the water (they couldn’t have just subsisted by wading in or eating what washed up, or at least this is what is implied), bodies started to change. Males are paid little attention by Morgan, and the warm relationship between mother and child takes center stage. While most of the hair on the body would be lost as an adaptation to water (an odd conclusion given that otters, seals, and sea lions all are covered in hair), the hair on the head would be allowed to grow long, the water babies being able to curl their fingers into it and stay close to mom for a nap when they got tired of exploring off on their own. Conversely, breast feeding would still have to take place on shore, but the upright posture of the females (acquired from so much time in the water) would require the baby to be held at an awkward position in which they could not reach their mother’s nipples. This was solved by developing larger “hemispherical” breasts to reach down to the infant, even though larger breasts may cause infants problems when they try to get their mouths on them to breast feed (if the breast is so large that the infant’s nose is covered by it, breathing and feeding becomes difficult).

In searching for an aquatic example of such a striking characteristic, Morgan turns to the Florida manatee and other sirenians, many who have seen females with young noting the presence of “breasts” on the aquatic mammals. Interestingly enough, however, the manatee shares it’s ancestry with living elephants, the females of which also exhibit some rather sizable swellings when lactating. Robert Sapolsky, in his book A Primate’s Memoir, describes seeing such an unexpected shape on the chest of a female elephant for the first time;

Did you know that female elephants have breasts? I do not mean rows of teats, a mama elephant lying on her side with dozens of little piglet elephants nursing with their eyes still closed. I mean breasts, two huge voluptuous billowy mounds, complete with cleavage. I bet you had no idea, did you? Nor did I – it is a subject rarely broached in our public schools. I’m out in the bush that first month, armed with binoculars and stopwatch and notepad, spending the days carefully watching baboons mating left and right. And then, suddenly, some pachyderms come cruising past, and I see some elephant with these, well, breasts. And the natural first reaction is to think, Oh, great, I’m such a horny lascivious pathetic adolescent that after a mere month of isolation in the bush I’ve already cracked, I’m hallucinating breasts the size of Volkswagens on the elephants. Horrors, to have one’s psychotic break occur so soon, and to have it take the form of a puerile sexual obsession many embarrassing steps below gawking at National Geographic nudies. I was greatly relieved to eventually discover that the elephant’s breasts were real, that I was not having some Marlin Perkins wet dream.

It should be noted, however, that Morgan attributes an aquatic origin to elephants as well, primarily based upon their ability to shed tears (and therefore salt), as well as the ability of living Indian elephants to swim long distances in the ocean. Such considerations are a side trip from the main thrust of her argument, and no detail is given as to when, where, why, or how elephants arose from a water-dwelling species, only that a few characteristics in living animals point to an Aquatic Pachyderm Hypothesis.

Going back to the AHH, given about 10 million years in the water Morgan’s ape is substantially different than the one that was first driven into the waves by a predator. Referring to her as “Mrs. Australopithecus,” Morgan paints the following portrait (the artistic manifestation of which is found on the cover of the book);

So our hominid has a nose. I have no doubt that she also had fleshy nostrils, but considerable doubt that they evolved to make sex sexier for her mate. I think she was by no means the simian, cadaverous, lipless creature that artists sometimes reconstruct by covering her dug-up skull with a tightly fitting layer of hairy skin. The layer of fat which was rounding out her arms and legs and adding bulk to her breasts was also filling out her cheeks, and her nostrils, and her earlobes, and everting her lips… We would not have accounted her beautiful, with her low forehead and prognathous jaw, but the chances are that she was a chubby little creature with several superficial features resembling our own more nearly than they resembled any ape’s. And as for the expressions that flitted across that prehistoric countenance, her millions of years in the water had certainly left their mark on those also.

This is quite a different picture of “Lucy” than is often seen, but is there anything to it? Part of the advantage of the AAH is that Morgan doesn’t specify her ideas down to a scientific level, allowing her to poetically play with her ideas in any way she wishes, the female becoming more beautiful while the men continue to try and kill dugongs with rocks. This type of feminist reaction to the “Man the Hunter” narrative is the main connective feature throughout the book, and Morgan’s writing is far more concerned with the more graceful and beautiful evolution of woman, with sex ultimately bringing “sin” into the Garden.

In Morgan’s story, the genitals of the ancestral females went from facing backwards (making rear-mounting positions by the male easy) to facing downwards, a position that Morgan insists will not work for males, face-to-face mating being adopted as a must. Morgan’s reasoning for the change is that aquatic animals often undergo this type of genital shift (cetaceans are her primary example), but she generally ignores why the genitals should be shifted in the first place. In terms of cetaceans, the ancestors likely had their anal-genital openings in the position typical for quadrapeds; facing backwards at the location of the pelvis just under the tail, usually being at the most distal end of the body. As they evolved, the archaeocetes lost their hind limbs and their spines elongated, being the main source of propulsion, so rather than keep moving backwards with the spine the genitals stayed in the pelvic region “settled” on the ventral side of the body; where else they would have gone, I do not know. Given this morphological necessity, face-to-face mating became the only way cetaceans could copulate. Seals and sea lions, on the other hand, still have their anal-genital openings near the distal most parts of their bodies because that is where the pelvis is and there was no need to change mating styles, and males still mount females from behind. Even beyond such considerations, I do not see how the rear-mount strategy can be dogmatically ruled out, and I have a feeling that because such a position is considered “kinky” by some it was essentially ruled not to have happened. In fact, the retention of rear-mounting with the shift in female genitals could help explain elongation of the penis in males (they’d have to extend a bit farther), although this matter is far from settled. Curiously, Morgan generally ignores the bonobo and it’s face-to-face mating habits, even in her later books. She’s clearly aware of these apes (she does mention them and one graces the cover of The Descent of the Child), but they are conspicuously absent from discussions about sex.

Still, if we are to follow Morgan’s model, the apes would have to switch from mating using a rear-mount position to face-to-face (the males, we are told, couldn’t penetrate any other way), such a position causing much trauma for females. Males wouldn’t know how to calm the female for a face-to-face encounter, and it essentially led to either rape or an unfruitful attempt to mate. Morgan describes such a scene;

The primate was a totally different shape. Her new aquatic streamlining had been unable to prevent her becoming lumpy in the middle, and as a littoral biped her legs were developing in the opposite direction from the seal’s – they were becoming not smaller and thinner but farther apart, but longer and thicker and closer together. The seal’s solution was impossible for the aquatic apes. Their dilemma was unique.

So we left her on her back, kicking and struggling and frightened out of her tiny anthropoid mind, with her mate beginning to get irritated. When she saw him snarl and bare his canines she was finally convinced that he wanted her for dinner, and that her last hour had come. Further resistance was useless. She stopped fighting and signaled her submission, defeat, and appeasement as strongly as she could with so little room for maneuver.

Immediately, the incident was over. The male was a properly programmed animal, and it was impossible for him to go on clobbering a member of his own species that was giving clear indications that it had stopped fighting back. He moved a little way off, wearing a puzzled expression. He had thought for a moment that he was on to a good idea, but obviously there was a snag to it.

Such events removed us from our Eden along the shores, males taking up hunting on the plains soon after the eviction. Rains that quenched the African drought allowed the apes to leave the habitat that they had become so accustomed to (it seems like the males led the charge, being sick of their prolonged day at the beach), moving on to evolve in ways that fit the scientific orthodoxy of the times a bit more closely. Even so, Morgan suggests that women have retained the peace, beauty, and grace of their aquatic origins while males are more shaped by violence and hunting, her parting words being;

He is the most miraculous of all the creatures that God ever made or the earth ever spawned. All we need to do is hold out our loving arms to him and say: “Come on in, the water’s lovely.”

Oddly enough, such arguments seem more specific and in-depth than those in Morgan’s later works The Descent of the Child (1995) and The Scars of Evolution (1990). The Descent of the Child can largely be ignored, being that it’s primary focus is on doing for human babies (from conception through early childhood) what The Descent of Woman did for women, all-in-all being a string of facts presented to the reader in an easy-to-digest manner but without much further discussion. In covering past evolution, the “savanna hypothesis” and “man the hunter” are both alluded to or pointed out to be wrong, although no rigorous refutation is made. Instead the reader is referred to the earlier The Scars of Evolution for the “scientific” argument, but Morgan’s earlier poetry contains far more detail than the 1990 work. I breezed through the 178 pages of the book easily enough, but there was little positive evidence within it’s pages for the AAH. Certain physiological systems were pinpointed and deemed to be of aquatic origin since Morgan deemed no other hypotheses to be adequate (which, of course, assumes that all possibilities have been discovered and have received proper consideration).

I actually would love to write up a longer discussion of The Scars of Evolution but there is surprisingly little actual AAH evidence to be considered, and Morgan even makes some fairly basic mistakes about fossil preservation. Early on in the book she writes;

So if the prospecting had started in the north [of the Rift Valley] and worked down, popular illustrations of groups of Australopithecus would have shown them reclining under a shady tree at the water’s edge, living perhaps on fruit and greenery and fish. Instead, they are depicted as shaggy creatures trekking through parched grass and a scatter of stunted thorn bushes, turning to scavenging and hunting to supplement their diet.

This conclusion comes from Morgan’s assertion that some specimens of Australopithecus are found associated with fossils like crocodile remains and turtle eggs, suggesting an aquatic habitat. This largely ignores taphonomy, however, and an animal that dies in or near water being much more likely to be preserved and fossilized than one that drops out on the plains, the body undoubtedly being ripped apart by scavengers and leaving little or nothing to the fossil record. Most of the rest of the book covers material already mentioned in The Descent of Woman, like the fallacious notion that pheromones are essentially nonexistent or non-influential in humans because we went through an aquatic phase of evolution where scent wouldn’t have counted for much. Also curious is one of Morgan’s final statements about how evolution works, especially in regards to water. Rather than gaining specializations mentioned in so many of her works (i.e. the ability to cry and remove saline from the body, nostrils with possible flaps to keep water out, enlargement of the female breasts), a kind of de-volution of our ancestors is favored;

Conceivably, a species finding itself in a radically new environment (such as water) begins to shed the more advanced features which fitted it for its old environment. It back-tracks to a more unspecialized foetus-like form, before re-adapting to the new habitat. If that were the case, then our own ancestors, having moved from the land to the water and subsequently from water to land, would have been subjected to an impetus towards neoteny on two successive occasions. It would explain why in our case the trend was unusually powerful.

In all, Morgan’s work seem to be lacking of any rigorous research or hypotheses, and it led me to wonder why the AAH will simply not go away. Perhaps some of it is the mental appeal and the common error of linking correlation in evolutionary convergence to causation, working backwards to whatever ideal we hold most dear. Even if I’m incorrect as far as social motivation goes, the AAH has shown up in the scientific literature in the past few years, and it’s primary advocate seems to be Marc Verhaegen. Although the majority of his papers seem to be currently unavailable online, there is no name that more frequently appears in terms of AAH literature in scientific journals, giving the hypothesis some visibility (and credibility, as far as AAH advocates may be concerned). Some of the papers published on the AAH I could find are;

Bender R, Verhaegen M, & Oser N. “Acquisition of human bipedal gait from the viewpoint of the aquatic ape theory” Anthropol Anz. 1997 Mar;55(1):1-14.

Cunnane, S.C. “The Aquatic Ape Theory reconsidered” Medical Hypotheses Volume 6, Issue 1, January 1980, Pages 49-58

Ellis, D.V. “Wetlands or aquatic ape? Availability of food resources.” Nutr Health. 1993;9(3):205-17.

Rhŷs Evans, PH. “The paranasal sinuses and other enigmas: an aquatic evolutionary theory” J Laryngol Otol. 1992 Mar;106(3):214-25.

Vaneechoutte, M. ” Report of the Symposium ‘Water and Human Evolution’, Gent, Belgium, April 30th 1999” Human Evolution. 2000 Volume 15, Numbers 3-4

Verhaegen, M.J.B., Puech, P.F., & Munro, S. “Aquarboreal ancestors?” Trends in ecology & evolution (Amsterdam). 2002 Vol. 17, Issue 5, page 212

Verhaegen, M.J.B. and Puech, P.F. “Hominid lifestyle and diet reconsidered: paleo-environmental and comparative data” Human Evolution. 2000 Volume 15, Numbers 3-4

Verhaegen, M.J.B. “The Aquatic Ape Theory and some common diseases”. Medical Hypotheses
Volume 24, Issue 3, November 1987, Pages 293-299

Verhaegen, M.J.B. “The Aquatic Ape Theory: Evidence and a possible scenario”
Medical Hypotheses Volume 16, Issue 1, January 1985, Pages 17-32

Verhaegen, M.J.B. “Aquatic ape theory and fossil hominids” Medical Hypotheses Volume 35, Issue 2, June 1991, Pages 108-114

Verhaegen, M.J.B. “Aquatic ape theory, speech origins, and brain differences with apes and monkeys” Medical Hypotheses Volume 44, Issue 5, May 1995, Pages 409-413

[And for an opposing view see Langdon, J.H. “Umbrella hypotheses and parsimony in human evolution: a critique of the Aquatic Ape Hypothesis” Human Evolution. 1997, Volume 33, Number 4, pp. 479-494(16)]

As is immediately apparent, the great majority of the papers have appeared in one journal (Medical Hypotheses) and can be attributed to one author, Verhaegen. Judging from what I was able to find, many of the arguments that Verhaegen employs are very similar to those of Morgan, working backwards from somewhat contested or enigmatic human features to an aquatic origin to the exclusion of other hypotheses. Where Verhaegen differs, however, is that his aquatic hypothesis is far more broad than that of Hardy or Morgan. While Morgan implied that the aquatic apes were an isolated group that ended up leading to man (what happened to populations elsewhere is never spelled out), Verhaegen suggests that the last common ancestor of living gorillas, chimpanzees, bonobos, and humans was at least semi-bipedal and semi-aquatic, likely living in a habitat like a mangrove swamp. From the paper “Aquaborel ancestors?”;

A vertical posture and an ability to climb with the arms raised above the head could have helped a wading primate to enter or leave the water by grasping overhanging branches or waterside vegetation, and to grasp fruits above the water. Body enlargement and tail reduction would hinder agile arborealism, whereas a larger body is more easily supported in water and helps reduce heat loss (explaining why aquatic mammals are larger than related terrestrial forms). Tails would be of little use for a wading and/or swimming primate and would cause both drag and heat loss.

Thus Verhaegen attempts to separate New and Old World monkeys from apes by making the ancestors of all living apes at least partially water-bound, standing up to wade through water. Ultimately humans would have stayed in the pool while gorillas and chimpanzees got out, although gorillas, chimpanzees, and bonobos do not seem to show the same signs of being adapted to water that are often associated with humans under the AAH. Of further note is the fact that living primates like baboons, macaques, and proboscis monkeys have been known to swim and stand upright in water, although none seem to show signs of becoming exclusively adapted to an aquatic lifestyle. In the recent BBC series Planet Earth, baboons of the Okovango Delta in Botswana were shown wading through the water;

The baboons are not especially comfortable in the water, just as many other animals in the delta like cheetahs and lions don’t especially like crossing the waterways. Indeed, crocodiles are the primary danger in the water, and many animals seem to know of the threat all too well (but must cross from time to time, anyway). Such modes of moving through water, also seen in chimpanzees (see the final episode of the BBC’s Life of Mammals, entitled “Food for Thought”), seem to constitute the “weak” version of the AAH, and isn’t entirely unreasonable in explain possible motivation to become bipedal. It does require a certain ecology, however, (i.e. flooded plains, a swamp, shallow mangrove forest, lagoon, etc.) and has little explaining power out of such a context. Still, there are even more aquatic primates that were also featured in Planet Earth; the crab-eating macaques.

If we’re looking for a model of what an aquatic ape would look and act like, surely these monkeys would be it, and Morgan does note some of the aquatic habits of macaques (especially the behavior of washing food in water, as seen in Japanese Macaques). The problem is that macaques are monkeys, not apes, although they seem to get along in the water just fine. Unfortunately the adaptation of these primates to water is going to be slow and take many generations, but the study of these animals could give us some clues as to what the AAH can and cannot explain, although it seems that many of the features explained by the AAH don’t fit with what we see in the macaques. Looking at the underwater behavior, it would seem that the monkeys would be adapted to swim in a matter similar to that of quadrupeds rather than to start wading in, becoming bipedal, and then doing a breast-stroke. Indeed, the video shows that becoming bipedal is not a necessary precursor to being able to swim or becoming semi-aquatic, and it is quite possible (even probable) that primates could abandon the upright stage altogether. Standing upright seems to be generally uncomfortable for many primates, and it’s hard to see how primates introduced to a fruitful aquatic habitat would want to stand up before just jumping in if there was really nothing to fear in the waters. Even in the weak version of the AAH, it is hard to see how standing upright while crossing a river would have selected for bipedalism as it seems that many primates are capable of doing it over short periods and it does not hold any strong advantage that would relate to mating success or overall survival. Unless the hypothetical apes lived in an area constantly flooded, requiring them to stand up much of the time, it is difficult for me to imagine how water could have helped to select for an upright posture.

The overall problem I have with Morgan’s hypothesis about apes becoming almost exclusively aquatic is that it forces us to make a choice of one habitat or another. Mentions are made of Proboscis Monkeys and Macaques enjoying a swim, washing off food, or living near water, but they don’t seem to be bent on the same path as the one Morgan proposes. Organisms certainly are plastic, and they don’t rigidly abide by the “rules” set down by those that describe them as to where to live, what to eat, and how to act properly. In fact, it seems more reasonable to me that primates past and present would take advantage of an aquatic resource if readily available, but still maintain their terrestrial life unless they were so isolated that they had no choice for food other than the water. Time will tell if some of today’s semi-aquatic primates ever become more fully at home in the water, but I see no reason to believe that our ancestors decided to take a prolonged summer vacation on the beach, proceeding in a way that just so happened to explain everything neatly (if un-parsimoniously).

The AAH hinges on apes willingly going into the water for safety from predators, but this is only a Just-So story without the details. It also ignores the fact that the water can be almost as dangerous, if not moreso, than the land, and there are predators in the water just as there are in terrestrial habitats, not to mention rip-tides and other problems inherent to the ocean itself. While Morgan, in The Descent of Woman, states that newborns could be left to paddle about on their own while mom went about her own business, such maternal inattention doesn’t seem like it would be especially effective in making sure that young made it to adulthood. Only the calmest, most sheltered, and safe of lagoons would have allowed for this. If the AAH is to be taken seriously in whatever form, it is going to require rigorous ecological study, and so far it seems that it relies far more on post hoc arguments than actual evidence.

While Jim Moore has already done a fantastic job dismantling the various problems with the AAH, I hope I have helped to illuminate the overall lack of evidence for the idea. As an idea it’s not a bad one, but it seems to have never gone beyond hypothetical situations and Just-So stories, and most of the ideas associated with the AAH seem to be criticisms of other hypotheses, therefore leaving the AAH as the only alternative. While I can certainly appreciate the frustration Morgan and others must have felt (and even still feel) towards a male-dominated field in science and consideration being mainly given to the strong, archetypal male, I feel that the AAH is taking things too far in the other extreme. It is hard to ignore the feminist underpinnings of Morgan’s writing and the overall disregard for the big picture in order to bring women and children into closer focus. Combating a hypothesis you don’t like with an equally narrow one, just reversed, is not the way to bring greater understanding of our evolutionary history, and given that hominids and apes are so close to us, it’s easy to fall into trapping of preference. Being that I am no expert on the matter, however, I will close with T.H. Huxley’s final words from his work Man’s Place in Nature, as they seem to resonate with the big questions about our origins that remain unknown;

Where, then, must we look for primeval man? Was the oldest Homo sapiens pliocene or miocene, or yet more ancient? In still older strata do the fossilized bones of an Ape more anthropoid, or a Man more pithecoid, than any yet known await the researches of some unborn paleontologist?

Time will show. But, in the meanwhile, if any form of the doctrine of progressive development is correct, we must extend by long epochs of the most liberal estimate that has yet been made of the antiquity of Man.

Pictures from Petsitting and the Delaware Water Gap

These are a bit long in coming, but here are some photos taken while petsitting a few weeks ago and during the trip to the NJ side of the Delaware Water Gap. Unfortunately I’m not too familiar with fungi so I can’t say I know what many of the species pictured are, but many of them were impressive all the same.

This spider was busy building a web outside the house I was staying at a few weeks ago. I’ve never seen an abdomen on a spider like this one has.

I usually only see Blue Jays during the winter (or at least only remember seeing them during winter), but this one stopped by the bird feeder.

A Cardinal pair also came by, although they were more skiddish and difficult to photograph.

Eastern Goldfinch and Cedar Waxwing were the most common visitors to the feeder, however.

A Turkey Vulture also circled overhead for a while, although it didn’t find anything interesting in the yard.

White-Tailed Deer also came by many times during the day, although the amount of brush and shrubs made them a little hard to photograph.

The second round of summer fawns also came by in the mornings and evenings, usually.

A very large Katydid with a color pattern I hadn’t seen before also paid us a visit, albeid inside the house.

Now on to the photos from the hike along the Appalachian Trail to Sunfish Pond;

Early on we came across these two fighting harvestmen (“daddy long legs”).

Being that the ground was relatively damp, there was fascinating fungi everywhere. The last two shots are among my favorites, and could the “S” on that last one be intelligently designed? (If you are new here, sprinkle the last bit of that sentence liberally with sarcasm)

Once we got up to Sunfish Pond, we were greeted by scores of Bullfrog and Leopard Frog young, which hopped, almost in unison, back into the shallows.

Toads were also present over the entirety of the hike. We counted at least 20 over the 10 miles.

We also saw two Five-Lined Skink (thanks for the correction, Lars) on a log and tree near the pond, the one on the tree have a brilliant blue tail.

The area that we sat down to lunch at was absolutely full of life as well, from fish and frogs to a small snake that was getting ready to shed.

In all, I’ve been able to get more photographs of NJ wildlife this summer than I have in previous years, and I hope that next year I’ll be able to get some better pictures overall.

Not your average bovid; the Saiga

A Mhorr Gazelle, a subspecies of Dama Gazelle, on a cold February morning at the Philadelphia Zoo

A male Gerenuk, taken in the summer of 2006 at Disney’s Animal Kingdom Park

When I think about antelope, something like a Thompson’s Gazelle, Gerenuk, or Dama Gazelle most immediately comes to mind, countless nature documentaries featuring the bovids in such numbers that they are almost easy to ignore as scenery (or as merely prey for the big cats and other carnivores). There is at least one antelope, however, that would make anyone do a double take; the Saiga (Saiga tatarica).

A Saiga, Saiga tatarica, from Wikipedia.

By all accounts the Saiga is quite distinct from its cousins on the African plains. Preferring steppe, semi-desert, and desert habitats, the Saiga is known from Eastern Europe through Mongolia, their conspicuous nose warming air coming into the body in the winter and filtering out dust during the summer. While this aspect of the Saiga’s adaptation is certainly fascinating, it is the horns of the animals that have brought it the most attention, however, as well as the most trouble. As I noted in a post about antlers, horns, and sexual selection, research has shown that males and females of bovid species (which includes antelope) that use their horns as defense against predators both have horns, the horns prevented from getting too gaudy in males because they need to retain their defensive function. In the case of the Saiga, only the males carry horns, and this allows for the horns to become relatively large, and this has greatly contributed to the decline of the species.

A male Saiga, from Sokolov, Mammalian Species, No. 38, Saiga tatarica (May 2, 1974), pp. 1-4

One of the primary problems that conservation officials face in Africa and Asia today is the poaching of animals for certain parts of their anatomy for use in Traditional Chinese Medicine (TCM), or other ritual aspects of a particular culture. To understand why the Saiga is targeted for its horns, however, we need to first look at the problems with rhinoceros poaching. Rhinoceros horns have long been treasured by various cultures, young men in Yemen traditionally carry a dagger for defense called a jambia, and there is no better material for the handle of this weapon than rhino horn. Initially, the amount of rhino horn taken was mitigated by the poverty in the region, only the most affluent families being able to afford a jambia with a handle made of genuine rhino horn. As oil prices went up, though, Yemen (among other countries) was flooded with income from the sale of fossil fuels, and the demand for rhino-horn-handled jambias skyrocketed.

Jambias are not the only reason rhinos have been slaughtered. In addition to various ornamental products, rhino horn is valuable in TCM as a fever-reducer. When actually tested it seemed to reduce fever in rats, and Saiga horn had about the same efficacy, although author Richard Ellis rightly notes in his book No Turning Back that “Asprin and ibuprofen, for which no animals have to die, would probably work just as well.” In India rhino horn was used as an aphrodisiac, but expensive prices caused practitioners to stop prescribing it. These pressures, along with others, ultimately drove the rhino population down so low that horns were rare even within illegal trade, and demand kept going up. A substitute had to be found.

The horns of the Saiga, the species only recently recovered from population declines in the early to mid-20th century, were actively endorsed by the World Wildlife Fund as a substitute for rhino horn (also see here and here. Strangely, while the WWF is one of the groups responsible for the crash of Saiga populations, they make no mention of their actions on their website, their efforts of helping the Saiga since 1994 being extolled rather than their responsibility for the near-eradication of the animals. Nevertheless, the political destabilization of Russia and other areas in which the Saiga lived made regulation near impossible, and while some researchers claim that the plains were once “blackened” with Saiga, there are probably less than 50,000 of the animals left throughout their entire range (the Mongolian Saiga, a subspecies, being reported as having a population of less than 1,300 in a 1999 paper by Lushchekina et al. in the journal Oryx).

Indeed, the tale of the Saiga is one of the greatest tales of ecological mismanagement in history, and its implications can still be felt today. Ellis, again in No Turning Back, tells us how it is unlikely that Saiga were as well-established by the time the WWF endorsed hunting Saiga as some conversationalists claim;

By the time of the Soviet Revolution, there were only a few thousand saigas left. To forestall their total eradication, the Soviets protected them in Europe in 1919 and in Soviet Central Asia in 1924. In the 1950s commercial harvests of saigas by local groups began.

Strangely, Saiga was not especially well-established in TCM until recently. Earlier texts like the 1597 text Chinese Materia Medica having no mention of the Saiga, but the 1989 Rare Chinese Materia Medica, and the ground-up horn of the male Saiga can be used to;

…check hyperactivity of the liver and relieve convulsion, treat the up-stirred liver wind, infantile convulsion and epilepsy; calm the liver and suppress hyperactivity of the liver-yang; it is efficacious in the treatment of dazzle and vertigo due to hyperactivity of the liver-yang; it improves acuity of vision, cures headache and conjuctival congestion; clears away heat and toxic material; and can be used to treat unconsciousness, delirium, and mania in the course of epidemic febrile disease.

So it does seem that the Saiga has been a more recent alternative to other traditional remedies, gaining the status of an “ancient treatment” only towards the end of the last century (although I am no expert on TCM and there could be earlier references than the ones mentioned by Ellis). As you probably have guessed by now, though, the major problem in Saiga conservation is that it is the males who are constantly targeted, much of remaining populations being females. This greatly reduces the amount of offspring that are likely to be produced, as well as marking a steep drop-off in genetic diversity and possibly even fitness in the species, the Saiga going through two near-extinctions in less than 100 years.

Male Saiga skull. Note the large shelf in front of the large nasal opening. From Sokolov, Mammalian Species, No. 38, Saiga tatarica (May 2, 1974), pp. 1-4

While the earlier near-extinction of the Saiga typically gets less attention than the more recent run on their horns, I feel that it echoes recent policies in the United States to remove wolves and other animals from protection; when stocks seem to approach levels that are barely adequate, many want to open up hunting again when the animals should actually be left alone to reestablish themselves and their genetic diversity. While some are optimistic about the recovery of Saiga, I do not share the same hope that they do. Even if raw numbers of Saiga continue to rise, I worry that the decreased genetic diversity will make them more susceptible genetic problems caused by inbreeding and disease epidemics, one “bad day” in an ecological sense being able to wipe the species out.

I would be remiss, however, if I attributed the problems of Saiga conservation entirely to hunting. As I just suggested, disease and parasites can be an important factor in terms of whether populations die off or not, something that can not be planned for by merely making sure there is more of the animals next season. In 1992, Dukes, et al. published a study that showed that paratuberculosis (or Johne’s Disease) was transmittable from domestic sheep to captive Saiga and back again. Even more recently (Morgan et al, 2005) it has been found that Saiga pick up many of the parasitic worms that also infect grazing livestock along their range, both suffering from the parasites and also allowing them to spread further. Global climate change may also adversely affect the Saiga, the Saiga depending on local climatic cues to dictate their migrations. Heavier snowfall/harsher drought may cause them to have to move to new habitats or be decimated, and at present it is unknown whether they’d be able to cope with ecological changes associated with the current warming trend.

In all, things are not looking too good for the Saiga. While there has been a ban on the trade of products made from them and conservation agencies are trying hard to preserve this species, I believe that it not exists in a weakened state, which (despite population size) will make it more susceptible to extinction. Given enough time the species may recover fully, but in my own view this has “heath hen” written all over it, a species that seemed to recover until disease, predation, and weather changes were too much for the remaining birds in New England to handle. I am not suggesting that we say the Saiga is a lost cause; far from it. What I am recommending is that we actually learn something from the terribly mistakes we’ve made with this species, and stop thinking that merely because populations increase it somehow equals immediate species stability.

Hoofnagle gets some things right, others wrong on Ecology

Over on Denialism Blog, Mark Hoofnagle as a rather, shall I say, “interesting” piece up on the interaction between man and nature, and he picks none other than Springfield’s Mr. Burns to show us what our guiding ecological principles should be;

“Oh, so Mother Nature needs a favour? Well, maybe she should have thought of that when she was besetting us with droughts and floods and poison monkeys. Nature started the fight for survival and she wants to quit because she’s losing? Well, I say hard cheese!”

[From one of my favorite episodes, “The Old Man and the Lisa“]

Burn’s position is patently short-sighted and foolish (and Hoofnagle admits it’s inclusion is somewhat facetious, but it’s not too far off the mark from what Mark proposes in the rest of his piece. He writes;

Nature is not our friend, nor have we advanced our species by living in harmony with it. We have survived, and tacked decades onto our lives by bending nature to our will. Nature is trying to kill us. All the time. Bacteria, and parasites and viruses oh my, they’re out to get us. We’re not buddy-buddy with nature, we’re in competition with it for our very survival, all the time. Further there is this nonsense that messing with nature is somehow a bad thing, or frequently unsuccessful. This view can only be held by people who seem to have forgotten all the progress we’ve made in the last couple of millennia.

Strong words, indeed. While I’m not one of the Disney-fied, harmonious “Circle of Life” view of nature (hippo carnivory and cannibalism can show us that, at least), this passage is little more than a straw man argument. “Nature” is not out to get us any more than it is helping us; there are no microbes or “poison monkeys” conspiring to do us in. There is competition and cooperation, each individual organism trying to meet its own needs, but there is no great war in nature where one species or “realm” of nature consciously seeks to destroy another (that is, except for humans). Charles Darwin understood the interplay between competition and the “cooperation” that can arise from it in his famous “tangled bank” passage;

It is interesting to contemplate a tangled bank, clothed with many plants of many kinds, with birds singing on the bushes, with various insects flitting about, and with worms crawling through the damp earth, and to reflect that these elaborately constructed forms, so different from each other, and dependent upon each other in so complex a manner, have all been produced by laws acting around us.

Hoofnagle’s mention of bacteria in solely a bad light is surprising as well; not all bacteria are bad for us. Probiotic bacteria found in our stomachs can be beneficial in binds to sites so that other, harmful bacteria can’t get enough elbow room to get settled in, and it’s been shown that some may even help combat lactose intolerance, diarrhea, and other problems. Even stepping beyond this, the hereditary disease hemochromatosis, where iron is “locked away” in the body, may have even been beneficial to humans, making its hosts less susceptible to certain infections that would end life far earlier than hemochromatosis would.

I’m also a bit disturbed by Hoofnagle’s assertion that “Further there is this nonsense that messing with nature is somehow a bad thing, or frequently unsuccessful. This view can only be held by people who seem to have forgotten all the progress we’ve made in the last couple of millennia.” Our current problems with global climate change and pollution, among other things, show that we’re not too bright when it comes to ecology or our impact. While we might call ourselves a “success” in that we’ve subdued nature successfully, I think it’s downright foolish to use some subjective idea of “progress” as the measuring stick by which to see whether our actions have been “good” or “bad.” Hoofnagle then rattles off a list of “achievements” by which I wouldn’t be here typing this right now; from laying the foundation of a house to squashing a spider that’s crept indoors, humanity is defined by its need to “sanitize” nature in order to be comfortable.

In one sense, Hoofnagle is right; disease, parasites, predators, drought, famine, etc. have largely been eliminated in the Western World (although this doesn’t mean that they’re not still problems for the majority of the planet’s inhabitants), and doing so has allowed us to live the “comfy” lives that those lucky enough to be born in the right place in the right time to the right parents enjoy. Still, as I’ve just mentioned, much of the world does cope with all the “war machinations” of nature, and even moreso did our ancestors. Nature was never out to kill us; it was simply the situation we found ourselves in, and one way or another nature was suppressed so that we could live. So far, things have been going alright from the perspective of geological time. We’ve come very far very quickly, but I’m rather dubious as to the future. Sometimes I feel that we feel that we’re smarter than we actually are, that our growth has been too exponential, and sooner or later (be it a meteorite, a disease, climate change, or even all of those plus others) things aren’t going to be going so smoothly for us.

Hoofnagle closes with these sentiments;

One does not commune with an uncaring monster. You don’t live in harmony with that which tries to kill you, at best you strive for detentes. Outright war isn’t good, a victory in a war might just be ashes in our mouth. We need her to survive, we need her to be healthy, but we should never trust her, or let down our guard. Nature is out to get us, we survive by constantly waging our battles with her, and winning. Not being the type that thinks this planet belongs to us, or that we are particularly relevant in the grand scheme of things, there’s a good chance she might win the war. I’d like to avoid that if at all possible.

Again, this is a rather weak argument based upon the anthropomorphization of some amorphous “Mother Nature” which seeks to make life hard for us. It’s true that in order to spread across the globe as we have, the suppression of nature has been required, but there have always been trade-offs. Sure, you can eradicate certain pests or disease-vectors, but you’re going to have to inhale plenty of harmful fumes that’ll end up contributing to your death too, just a bit later. Sure, industrialization and the invention of the automobile has opened up untold wonders for us, but that isn’t going to mean much if climate change and disease radically change the planet, perhaps even causing a much higher death rate. I don’t believe in any notion of “progress” that does not have some sort of consequence somewhere, and while we have never lived in “harmony with nature,” being overly abusive to Hoofnagle’s “uncaring monster” isn’t going to get us very far either.

[Hat-tip to Pondering Pikaia for the story]

Curioser and curioser… AFM wants me to blog about DDT

I wonder if anyone else received a similar e-mail that just arrived in my inbox; Philip Coticelli of the group Africa Fighting Malaria mentions a new study that has come out in PLoS ONE called “A New Classification System for the Actions of IRS Chemicals Traditionally Used For Malaria Control.” The message I received featured the headline “DDT Highly Effective Against Resistant Mosquitoes: New study recommends using DDT to control #1 killer of African children: malaria,” so where the AFM stands on DDT use seems pretty clear.

I assume I was contacted because I mentioned how National Geographic made some errors in their cover story on Malaria over a month ago, although Bug Girl and Ed Darrell are probably much more capable bloggers to handle this one. Still, when I get the chance I’ll have a look at the article and write up some thoughts, despite that fact that even if DDT has proven effective against mosquitoes thought to be resistant, the efficacy of the product doesn’t assuage my concerns about its toxicity.