The hand of Homo naledi points to life before birth

Homo naledi is one of my favorite extinct humans, in part because its impressive fossil record provides rare insights into patterns and process of growth and development. When researchers began recovering naledi fossils from Rising Star Cave 10 years ago, one of the coolest finds was this nearly complete hand skeleton. The individual bones were still articulated practically as they were in life so we know which bones belong to which fingers, allowing us grasp how dextrous this ancient human was. And since finger proportions are established before birth during embryonic development, we can see if Homo naledi bodies were assembled in ways more like us or other apes.

The “Hand 1” skeleton of Homo naledi, adapted from a figure by Kivell and colleagues (2015). Left shows the palm-side view while the middle shows the back of the hand. The inset (b) shows many of the palm and finger bones as they were found in situ in Rising Star Cave.

In a paper hot off the press (here), I teamed up with Dr. Tracy Kivell to analyze finger lengths of Homo naledi from the perspective of developmental biology. On the one hand, repeating structures such as teeth or the bones of a finger must be coordinated in their development, and scientists way smarter than me have come up with mathematical models predicting the relative sizes of these structures (for instance, teeth, digits, and more). On the other hand, the relative lengths of the second and fourth digits (pointer and ring fingers, respectively) are influenced by exposure to sex hormones during a narrow window in embryonic development: this ‘digit ratio’ tends to differ between mammalian males and females, and between primate species with different social systems.

So, Tracy and I examined the lengths of the three bones within the second digit (PP2, IP2, DP2) and of the first segment of the second and fourth digits (2P:4P) in Homo naledi, compared to published data for living and fossil primates (here and here). What did we find out?

Summary of our paper showing the finger segments analyzed (left), and graphs of the main results (right). The position of Homo naledi is highlighted by the blue star in each graph.

The first graph above compares the relative length of the first and last segments of the pointer finger across humans, apes, and fossil species. The dashed line shows where the data points are predicted to fall based on a theoretical model of development. There is a general separation between humans and the apes reflecting the fact that humans have a relatively long distal segment, which is important for precise grips when manipulating small objects. Fossil apes from millions of years ago and the 4.4 million year old hominin Ardipithecus are more like apes, while Homo naledi and more recent hominins are more like modern humans. Because both humans and apes fall close to the model predictions, this means the theoretical model does a good job of explaining how fingers develop. Because humans and apes differ from one another, this suggests a subtle ‘tweak’ to embryonic development may underlie the evolution of a precision grip in the human lineage, and that it occurred between the appearance of Ardipithecus and Homo.

The second graph compares the ‘digit ratio’ of the pointer and ring fingers from a handful of fossils with published ratios for humans and the other apes. Importantly, the digit ratio is high in gibbons (Hylobates) which usually form monogamous pair bonds, while the great apes (Pongo, Gorilla, Pan) are characterized by greater aggression and mating competition and have correspondingly lower digit ratios. Ever the bad primates, humans fall in between these two extremes. Most fossil apes and hominins have digit ratios within the range of overlap between the ape and human ratios, but Homo naledi has the highest ratio of all fossil hominins known, just above the human average. It has previously been suggested that humans’ higher ratio compared to earlier hominins may result from natural selection favoring less aggression and more cooperation recently in our evolution. If we can really extrapolate from digit proportions to behavior, this could mean Homo naledi was also less aggressive. This is consistent with the absence of healed skull fractures in the vast cranial sample (such skull injuries are common in much of the rest of the human fossil record).

You can see the amazing articulated Homo naledi hand skeleton for yourself on Morphosource. Its completeness reveals how handy Homo naledi was 300,000 years ago, and it can even shed light on the evolution of growth and development (and possibly social behavior) in the human lineage.

What do capuchin stone tools tell us about human evolution?

A month ago at ESHE and now online in Nature, Proffitt and colleagues describe stone-on-stone smashing behavior among wild bearded capuchin monkeys (Sapajus libidinosus). The online paper includes a great video documenting the action; here’s a screenshot:

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Holding the rock with both hands just above head-level, the monkey prepares to crush its enemies. Which in this case are another rock stuck in a pile of more rocks.

In the fairly rare cases where non-human primates use stones, it’s for smashing nuts or something. But when these capuchins see a stone they don’t just see a smasher, they see a world of possibilities* – why use a rock to break a rock, when you could use it to break a heart? So this group of capuchins is unique in part because they’ve been documented to use stones for many purposes.

Now why on earth a monkey would use one rock to break another rock is anyone’s guess. In human evolution, the purpose was to break off small, sharp flakes that could be used to butcher animals or work plants. Proffitt et al. did observe small flakes being removed when capuchins pounded rocks, but the monkeys showed little interest in this debitage, other than using it to continue smashing stuff. More curiously, the monkeys frequently lick the rock after hammering at it:

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Mmm, rocks.

Proffitt et al. venture that maybe these monkeys are doing this to ingest lichens or trace elements like silicon. This hypothesis merits further investigation, but what’s clear is that these monkeys’ lithics differ from the hominin archaeological record wherein the express purpose of breaking rocks is to make flakes.

What’s striking to me (pun intended) is the relative size of the rocks. These monkeys that weigh only 2-3 kg are lifting and smashing stones that weigh about half a kilogram on average. Because these stones are fairly large given the monkeys’ body size, they have to be lifted with two hands and brought down on a surface, a “passive hammer” technique. The earliest-known tools made by hominins, from the 3.3 million year old Lomekwi site in Kenya, are also pretty big. Weighing 3 kg on average but topping at 15 kg, these earliest tools would have required the same knapping technique as is used by these little monkeys (Harmand et al., 2015).

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Left: Cover of Nature vol. 521 (7552). Right: Bearded capuchin letting a pebble know who’s boss (link).

Why the big stuff at first? Did the earliest hominin tool-makers lack the dexterity to make tools from the smaller rocks comprising the later Oldowan industry? These creative capuchins could lead to predictions about the hand/arm skeleton of the Lomekwian tool-makers (testable, of course, only with fortuitous fossil discoveries). Capuchins are noted for their manual dexterity (Truppa et al., 2016) and have a similar thumb-index finger ratio to humans and early hominins (Feix et al. 2015), although they differ from humans in the insertion of the opponens muscle and resultant mobility of the thumb (Aversi-Ferreira et al., 2014). Maybe these tech-smart monkeys can tell us more about the earliest human tool-makers’ bodies than their brains.

ResearchBlogging.orgReferences

Aversi-Ferreira RA, Souto Maior R, Aziz A, Ziermann JM, Nishijo H, Tomaz C, Tavares MC, & Aversi-Ferreira TA (2014). Anatomical analysis of thumb opponency movement in the capuchin monkey (Sapajus sp). PloS one, 9 (2) PMID: 24498307

Feix T, Kivell TL, Pouydebat E, & Dollar AM (2015). Estimating thumb-index finger precision grip and manipulation potential in extant and fossil primates. Journal of the Royal Society, Interface, 12 (106) PMID: 25878134

Harmand S, Lewis JE, Feibel CS, Lepre CJ, Prat S, Lenoble A, Boës X, Quinn RL, Brenet M, Arroyo A, Taylor N, Clément S, Daver G, Brugal JP, Leakey L, Mortlock RA, Wright JD, Lokorodi S, Kirwa C, Kent DV, & Roche H (2015). 3.3-million-year-old stone tools from Lomekwi 3, West Turkana, Kenya. Nature, 521 (7552), 310-5 PMID: 25993961

Proffitt, T., Luncz, L., Falótico, T., Ottoni, E., de la Torre, I., & Haslam, M. (2016). Wild monkeys flake stone tools Nature DOI: 10.1038/nature20112

Truppa V, Spinozzi G, Laganà T, Piano Mortari E, & Sabbatini G (2016). Versatile grasping ability in power-grip actions by tufted capuchin monkeys (Sapajus spp.). American Journal of Physical Anthropology, 159 (1), 63-72 PMID: 26301957

*well, at least four uses given by Proffitt et al.: mating display, aggression, food-crushing, and digging.