Osteology Everywhere: Ilium Nublar

June 30, 2015 / zcofran / 1 Comment

Jurassic Park is objectively the greatest film ever made, so I don’t need to explain why I recently watched it for the bajillionth time. Despite having seen this empirically excellent movie countless times, I finally noticed something I’d never seen before.

Hold on to your butts. What's that on the screen in front of Ray Arnold?

Hold on to your butts – what’s that on the screen in front of John Arnold? (image credit)

The film takes place on the fictitious island “Isla Nublar,” a map of which features prominently in the computer control room when s**t starts to go down. Here’s a clearer screenshot of one of Dennis Nedry‘s monitors:

Isla Nublar from the JP control room. Quiet, all of you! They’re approaching the tyrannosaur paddock…. (image credit)

It dawned on me that the inspiration for this island is none other than MLD 7, a juvenile Australopithecus africanus ilium from the Makapansgat site in South Africa:

Figure 1 from Dart, 1958. Left side is MLD 7 and right is MLD 25. Top row is the lateral view (from the side) and bottom row is the medial view (from the inside). These two hip bones are from the left side of the body (see the pelvis figure in this post). Note the prominent anterior inferior iliac spine on MLD 7, a quintessential feature of bipeds.

Isla Nublar is basically MLD 7 viewed at an angle so that appears relatively narrower from side to side:

MLD at a slightly oblique view (or stretched top to bottom) magically transforms into Isla Nublar.

MLD 7 at a slightly oblique view (or stretched top to bottom) magically transforms into Isla Nublar.

It’s rather remarkable that some of the most complete pelvic remains we have for australopithecines are two juveniles of similar developmental ages and sizes from the same site. In both, the iliac crest is not fused, and joints of the acetabulum (hip socket) hadn’t fused together yet. The immaturity of these two fossils matches what is seen prior to puberty in humans and chimpanzees. Berge (1998) also noted that MLD 7, serving as an archetype for juvenile Australopithecus, is similar in shape to juvenile humans, whereas adult Australopithecus (represented by Sts 14 and AL 288) are much flatter and wider side to side. Berge took this pattern of ontogenetic variation to match an ape-like pattern of ilium shape growth. This suggests a role of heterochrony in the evolution of human pelvic shape, or as Berge (1998: 451) put it, “Parallel change in pelvic shape between human ontogeny and hominid phylogeny.” In layman’s terms, ‘similar changes in both pelvic growth and pelvis evolution.’

eFfing #FossilFriday: Rekindling an old friend’s hip

June 12, 2015January 1, 2016 / zcofran / 2 Comments

Sorry for the crappy pun. Carol Ward and colleagues recently reported an associated hip joint, KNM-ER 5881, attributable to the genus Homo (1.9 million years old). Fossils coming from the same skeleton are pretty rare, but what’s more remarkable is that portions of this bone were discovered 29 years apart: a femur fragment was first found in 1980, and more of the femur and part of the ilium were found at the same location when scientists returned in 2009:

Figure 3 from Ward et al. 2015. A little distal to the hip, yes, but the pun still works. Views are, going clockwise starting at the top the top left, from above, from below, from the back, from the side, and from the front.

There’s also a partial ilium associated with the femur – that makes a pretty complete hip!

Figure 5 from Ward et al. shows the fossil. Jump for joy that it’s complete enough for us to tell it comes from the left side!

Despite how fragmentary the femur and ilium are, the researchers were able to estimate the diameter of the femur head and hip socket reliably. The hip joints are smaller than all Early Pleistocene Homo except for the Gona pelvis. Comparing ER 5881 the large contemporaneous KNM-ER 3228 hip bone, the authors found these two specimens to be more different in size than is usually seen between sexes of many primate species. The size difference best matches male-female differences in highly dimorphic species like gorillas.

Ward et al. find that the specimen generally looks like early Homo but that the inferred shape of the pelvic inlet is a little different from all other Early and Middle Pleistocene human fossils. The authors take this discrepancy to suggest that there was more than one “morphotype” (‘kind of shape’), and therefore possibly species, of Homo around 1.9 million years ago. While I wouldn’t just yet go so far as to say this anatomy is due to species differences, I do agree that KNM ER 5881 helps our understanding and appreciation of anatomical variation in our early ancestors. Like all great fossil discoveries, the more we find, the more we learn that we don’t know. Here’s to more Homo hips in the near future!

Ima Gona follow up on that last post

March 10, 2014March 10, 2014 / zcofran / 2 Comments

Last week, I discussed the implications of the Gona hominin pelvis for body size and body size variation in Homo erectus. One of the bajillion things I have been working on since this post is elaborating on this analysis to write up, so stay tuned for more developments!

Now, when we compared the gross size of the hip joint between fossil Homo and living apes (based on the femur head in most specimens but the acetabulum in Gona and a few other fossils), the range of variation in Homo-including-Gona was generally elevated above variation seen in all living great apes. This is impressive, since orangutans and gorillas show a great range of variation due sexual dimorphism (normal differences between females and males). However, I noted that the specimens I used were unsexed, and so the resampling strategy used to quantify variation within a species – randomly selecting two specimens and taking the ratio of the larger to smaller – probably underestimated sexual dimorphism.

Shortly after I posted this, Dr. Herman Pontzer twitterated me to point out he has made lots of skeletal data freely available on his website (a tremendous resource). The ape and human data I used for last week’s post did not have sexes (my colleague has since sent me that information), but Pontzer’s data are sexed (no, not “sext“). So, I modified and reran the original resampling analysis using the Pontzer data, and it nicely illustrates the difference between using a max/min vs. male/female ratio to compare variation:

Hip joint size variation in living African apes (left and right) compared with fossil humans (genus Homo older than 1 mya, center). Each plot is scaled to show the same y-axis range. On the left are ratios of max/min from resampled pairs from each species (sex not taken into account). On the right are ratios of male/female from resampled pairs from each species. The red stars on this plot are the medians for max/min ratios (the thick black bars in the left plot). The center plot shows ratios of Homo/Gona.

The left plot shows resampled ratios of max/min in humans, chimpanzees and gorillas, while the right shows ratios of male/female in these species. If no assumption is made about a specimen’s sex (left plot), it is possible to resample a pair of the same sex, and so it is likelier to sample two individuals similar in size. Note that the ratio of max/min can never be less than 1. However, if sex is taken into account (right plot), we see two key differences. First, because of size overlap between males and females in humans and chimpanzees, ratios can fall below 1. Adult gorilla males are much larger than females, and so the ratio is never as low as 1 (minimum=1.08). Second, in more dimorphic species, the male/female ratio is elevated above the max/min ratio (red stars in the right plot). In chimpanzees, the median male/female ratio is actually just barely lower than the median max/min ratio. If you want numbers: the median max/min ratios for humans, chimpanzees and gorillas are 1.09, 1.06 and 1.16, respectively. The corresponding median male/female ratios are 1.15, 1.06 and 1.25.

Regarding the fossils, if we assume that Gona is female and all other ≥1 mya Homo hips are male, the range of hip size variation can be found within the gorilla range, and less often in the human range.

But the story doesn’t end here. One thing I’ve considered for the full analysis (and as Pontzer also pointed out on Twitter) is that the relationship between hip joint size and body weight is not the same between humans and apes. As bipeds, we humans place all our upper body weight on our hips; apes aren’t bipedal and so relatively less of their weight is transmitted through this joint. As a result, human hip joint size increases faster with increasing body mass than it does in apes.

So for next installment in this fossil saga, I’ll consider body mass variation estimated from hip joint size. Based on known hip-body size relationships in humans vs. apes, we can predict that male/female variation in humans and fossil hominins will be relatively higher than the ratios presented here – will this put fossil Homo-includng-Gona outside the gorilla range of variation? Stay tuned to find out!

Gona … Gona … not Gona work here anymore more

March 4, 2014March 5, 2014 / zcofran / 2 Comments

The Gona pelvic remains (A-D), and the reconstructed complete pelvis (E-J), Fig. 2 in Simpson et al., 2008.

A few years ago, Scott Simpson and colleagues published some of the most complete fossil human hips (right). The fossils are from the Busidima geological formation in the Gona region of Ethiopia, dated to between 0.9-1.4 million years ago. (Back when I wasn’t the only author of this blog, my friend and colleague Caroline VanSickle wrote about it here)

Researchers attributed the pelvis to Homo erectus on the basis of its late geological age and a number of derived (Homo-like) features. In addition, the pelvis’s very small size indicated it probably belonged to a female. One implication of this fossil was that male and female H. erectus differed drastically in body size.

Christopher Ruff (2010) took issue with how small this specimen was, noting that its overall size is more similar to the small-bodied Australopithecus species. Using the size of the hip joint as a proxy for body mass, Ruff argued Gona’s small size would imply a profound amount of sexual dimorphism in H. erectus: much higher than if Gona is excluded from this species, and higher than in modern humans or other fossil humans. Ruff thus proposed an alternative hypothesis to marked sexual dimorphism, that the Gona pelvis may have belonged to an australopithecine.

Fig. 3 From Ruff's (2010) reply. Australopiths (and Orrorin) are squares and Homo are circles. Busidima's estimated femur head diameter is represented by the star and bar.

Fig. 3 From Ruff’s (2010) reply. Australopiths (and Orrorin) are squares and Homo are circles. Gona’s estimated femur head diameter is represented by the star and bar.

Now, Simpson & team replied to Ruff’s comments, providing a laundry list of reasons why this pelvis is H. erectus and not Australopithecus. They cite many anatomical features of the pelvis shared with Gona and Homo fossils, but not australopithecines. They also note that there are many other bones reflective of body size, that seem to suggest a substantial amount of size variation in Homo fossils, even those from a single site such as Dmanisi (Lordkipanadze et al., 2007).

Interestingly, neither of these parties compared the implied size variation with that of living apes. So I’ll do it! Now, I do not have any acetabulum data, but a friend lent me some femur head measurements for living great apes a few years ago. Gona is a pelvis and not a femur, but there are more fossil femora than hips. Because there’s a very high correlation between femur head and acetabulum size, Ruff estimated Gona’s femur head diameter to be 32.6 mm (95% confidence interval: 30.1-35.2; Simpson et al. initially estimated 35.1 mm based on a different dataset and method). To quantify size variation, we can compare ratios of larger femur heads divided by smaller ones. Now, this ratio quantifies inter-individual variation, but it will underestimate sexual dimorphism since I’m likely sampling some same-sex pairs that aren’t so different in size. But this is just a quick and dirty look. So, here’s a box plot of these ratios for Homo fossils, larger specimens divided by Gona’s estimated femur head size in different time periods:

Ratio of a fossil Homo femur head diameter (HD) divided by Busidima's HD. E Homo = early Pleistocene, Contemporaneous = WT 15000 and OH 28, MP = Middle Pleistocene Homo. White boxes are based on Ruff's Busidima HD estimate, green boxes are based on Simpson et al.'s estimate.

Ratios of fossil Homo femur head diameter (HD) divided by Busidima’s (Gona’s) HD. E Homo = early Pleistocene, Contemporaneous = WT 15000 and OH 28, MP = Middle Pleistocene Homo. White boxes are based on Ruff’s Gona HD estimate, green boxes are based on Simpson et al.’s larger estimate. Boxes include 50% quartiles and the thick lines within are sample medians.

Clearly, Gona is much smaller than most other fossil Homo hips, since ratios are never smaller than 1.14. Average body size increases over time in the Homo lineage, reflected in increasing ratios from left to right on the plot. Early Pleistocene Homo fossils are fairly small, including Dmanisi, hence the lower ratios than later time periods. Middle Pleistocene Homo (MP), represented by the most fossils, shows a large range of variation, but even the smallest is still 1.17 times larger than the largest estimate of Gona’s femur head size. To put this into context, here are those green ratios (assuming a larger size for Gona) compared with large/small ratios from resampled pairs of living apes and humans:

The fossil ratios of larger/smaller HD from above, compared with resampled ratios from unsexed living apes and humans. Boxes include the 50% quartiles, and the thick lines within are sample medians. **(05/03/14: This plot has been modified from the original version post, which only included the fossil ratios based on the smaller Gona estimate)

What we see for the extant apes and humans makes sense: humans and chimpanzees show smaller differences on average, whereas average differences between gorillas and orangutans are larger. This accords with patterns of sexual dimorphism in these species. **What this larger box plot shows is that if we accept Ruff’s smaller average estimate of Gona’s femur head size (white boxes), it is relatively rare to sample two living specimens so different in size as seen between Gona and other fossils. If we use Simpson et al.’s larger Gona size estimate, variation is still elevated above most living ape ratios. Only when Gona is compared with the generally-smaller, earlier Pleistocene fossils, does the estimated range of variation show decent overlap with living species. Even then, the overlap is still above the median values.

These results based on living species agree with Ruff’s concern, that including Gona in Homo erectus results in an unusually large range of variation in this species. Such a large size range isn’t necessarily impossible, but it would be surprising to see more variation than is common in gorillas and orangutans, where sexual size dimorphism is tremendous. Ruff suggested that the australopith-sized Gona pelvis may in fact be an australopith. This was initially deemed unlikely, in part because the fossil is well-dated to relatively late, 0.9-1.4 million years ago. However, Dominguez-Rodgrigo and colleauges (2013) recently reported a 1.34 mya Australopithecus boisei skeleton from Olduvai Gorge, so it is possible that australopiths persisted longer than we’ve got fossil evidence for, and Gona is one of the latest holdouts.

So many possible explanations. More clarity may come with further study of the fossils at hand, but chances are we won’t be able to eliminate any of these possibilities until we get more fossils. (also, the post title wasn’t a jab at the fossils or researchers, but rather a reference to the movie Office Space)

References

Dominguez-Rodrigo et al. 2013. First partial skeleton of a 1.33-million-year-old Paranthropus boisei from Bed II, Olduvai Gorge, Tanzania. PLoS One 8: e80347.

Ruff C. 2010. Body size and body shape in early hominins – implications of the Gona pelvis. Journal of Human Evolution 58: 166-178.

Simpson S et al. 2008. A female Homo erectus pelvis from Gona, Ethiopia. Science 322: 1089-1092.

Simpson S et al. In press. The female Homo pelvis from Gona: Response to Ruff (2010). Journal of Human Evolution. http://dx.doi.org/10.1016/j.jhevol.2013.12.004

Osteology everywhere: Pelvis has left the building

March 13, 2012August 4, 2013 / zcofran / 3 Comments

The vernal awakening has brought rain to Ann Arbor, and right on here on main campus I spotted the rain-splotched silhouette of an articulated human pelvis (left).

Check out those short and flaring iliac blades, and the shortness of the ischium. These features are associated with repositioning key muscles for walking and running on two feet, and are very unlike what is seen in the four-legged, suspensory climbing apes.

But just how ‘human’ are these features? The crushed pelvis of Oreopithecus bambolii, a ~8 million year old fossil ape from Italy, has somewhat human-like short ilia (left). This pelvis also has weak anterior inferior iliac spines (Rook et al. 1999), which anchor the hip/trunk flexor muscle rectus femoris, and are allegedly a developmental novelty seen only in hominids (Lovejoy et al. 2009). These traits have led some to claim that Oreopithecus was a hominid, or at least bipedal. Without getting into that debate, I’ll just say that seeing these ‘bipedal’ features in this late Miocene ape’s pelvis weakens the case that their presence in Ardipithecus ramidus indicates a unique connection between Ardi and later, true hominids like australopiths.

UPDATE: Check the comments for notes on the Ardi and Oreo fossils from someone who’s actually studied them (I myself have only seen pictures and read about them).

References
Lovejoy, C., Suwa, G., Spurlock, L., Asfaw, B., & White, T. (2009). The Pelvis and Femur of Ardipithecus ramidus: The Emergence of Upright Walking Science, 326 (5949), 71-71 DOI: 10.1126/science.1175831

Rook, L. (1999). Oreopithecus was a bipedal ape after all: Evidence from the iliac cancellous architecture Proceedings of the National Academy of Sciences, 96 (15), 8795-8799 DOI: 10.1073/pnas.96.15.8795

ARDIPITHECUS BEER!!!

November 12, 2011August 4, 2013 / zcofran / 4 Comments

I just made what what may be the most amazing discovery of the century at a local booze emporium. Dogfish Head brewing company makes a beer whose label is adorned with Jay Matternes’s reconstruction of an upright Ardipithecus ramidus. Note that the left foot grasps the earth with it’s ape-like big toe.

In a whimsical use of artistic license, whoever adopted this image added a curlicue pig’s tail. In animals with a tail, a number of caudal vertebrae continue off the set of fused vertebrae called the sacrum. Humans and other apes don’t have true tails but a coccyx, a small clump of tiny, fused vertebral segments. Our tail vestige may not help us hang onto trees like in Ateline monkeys, or sting our enemies like a scorpion, but the coccyx is still pretty important. In people this evolutionary memory of a tail anchors some muscles of the pelvic floor (including sphincter ani externus and levator ani), which are critical for the to control of our bowels.

Below is a close up of the Ardipithecus ramidus pelvis fossils (from White et al. 2009, fig. 3). No coccyx was discovered for Ardi, and little is said about the sacrum, other than that it’s merely broken piece of the end of the bone (Lovejoy et al. 2009). Nevertheless, I’m sure this end of sacrum would lead one to reject this artist’s hypothesis that Ardipithecus had a tail.

Had I been in charge of labeling at Dogfish Head, the beer would’ve been called “Party-pithecus” instead of “namaste,” and this label would’ve been slapped on some exotic IPA or porter instead of a wheat beer. Still pretty awesome, though.

Learn about Ardi and its pelvis
Lovejoy, C., Suwa, G., Spurlock, L., Asfaw, B., & White, T. (2009). The Pelvis and Femur of Ardipithecus ramidus: The Emergence of Upright Walking Science, 326 (5949), 71-71 DOI: 10.1126/science.1175831

White, T., Asfaw, B., Beyene, Y., Haile-Selassie, Y., Lovejoy, C., Suwa, G., & WoldeGabriel, G. (2009). Ardipithecus ramidus and the Paleobiology of Early Hominids Science, 326 (5949), 64-64 DOI: 10.1126/science.1175802

[insert clever quip about australopithecus hips]

September 19, 2011August 4, 2013 / zcofran / Leave a comment

A week and a half ago, Kibii and colleagues (2011) published reconstructions and re-analyses of two hips belonging to the 1.98 million-year old Australopithecus sediba. As with many fossil discoveries, these additions to the fossil record raise more questions than they answer. Unless the question was, “did A. sediba have a pelvis?” It did. Here’s a good summary from the paper itself:

Thus, Au. sediba is australopith-like in having a long superior pubic ramus and an anteriorly positioned and indistinctly developed iliac pillar…[and] Homo-like in having vertically oriented and sigmoid shaped iliac blades, more robust ilia, and a narrow tuberoacetabular sulcus…and the pubic body is upwardly rotated as in Homo. (p. 1410, emphases mine)

So far as I can tell, the main way the hips are ‘advanced’ toward a more human-like condition is that the iliac blades are more upright and sweep forward more than in earlier known hominid hips. Here’s the figure 2 from the paper (more sweet pics of the fossils are available here). NB that in both A. sediba hips much of the upper portions of the iliac blades are missing (reconstructed in white; this region is missing in lots of fossils), so it’s possible they were more flaring like the australopith in the center photo.

The authors’ bottom-line, take-home point is that the A. sediba pelvis has features traditionally associated with large-brained Homo – but belonged to a small-brained species (based solely on the ~430 cc MH1 endocast). They argue that this means that many of these unique pelvic features did not evolve in the context of birthing large-brained babies, as has often been thought. They state that these features are thus “most parsimoniously attributed to altered biomechanical demands on the pelvis in locomotion,” and suggest that this hypothetical locomotion was mostly bipedalism but with a good degree of climbing. Maybe, maybe not. This interpretation is consistent with the analysis of the A. sediba foot/ankle (Zipfel et al. 2011).

The weird mix of ancient (australopith-like) and newer (Homo-like) pelvic features in A. sediba really raises the question of how australopithecines moved around. More intriguing is that the A. sediba pelvis has different Homo-like features than the ~1 million year old Busidima pelvis (Simpson et al. 2008), which has been attributed to Homo erectus (largely in aspects of the iliac blades). This raises the question of whether A. sediba is really pertinent to the origins of the genus Homo, and whether the Busidima pelvis belongs to Homo erectus or a late-surviving robust australopithecus (e.g. boisei, Ruff 2010).

Also interesting is that the subpubic angle (in the pic above, the upside-down “V” created by the pubic bones just above the red labels) is pretty low in MH2. This is curious because modern human males and females differ in how large this angle is – females tend to have a large angle which contributes to an enlarged birth canal, whereas males have a low angle like MH2. But MH2 is considered female based on skeletal and dental size. This raises the additional questions of whether human-like sexual dimorphism had not evolved in hominids prior to 1.9 million years ago, and whether the sex of MH2 was accurately described.

Finally, though the authors did a great job comparing this pelvis with those from other hominids, I think a major, more comprehensive comparative review of hominid pelves is in order. How does the older A. afarensis hip from Woranso (Haile-Selassie et al. 2010) inform australopithecine pelvic evolution? What about the possibly-contemporary-maybe-later hip from the nearby site of Drimolen (Gommery et al. 2002)? Given the subadult status of the MH1 individual, it would be interesting to compare with the WT 15000 Homo erectus fossils, or A. africanus subadults from Makapansgat, to examine the evolution of pelvic growth.

Lots of interesting questions arise from these fascinating new fossils. “The more you know,” right?

References
Gommery, D. (2002). Description d’un bassin fragmentaire de Paranthropus robustus du site Plio-Pléistocène de Drimolen (Afrique du Sud)A fragmentary pelvis of Paranthropus robustus of the Plio-Pleistocene site of Drimolen (Republic of South Africa) Geobios, 35 (2), 265-281 DOI: 10.1016/S0016-6995(02)00022-0

Haile-Selassie Y, Latimer BM, Alene M, Deino AL, Gibert L, Melillo SM, Saylor BZ, Scott GR, & Lovejoy CO (2010). An early Australopithecus afarensis postcranium from Woranso-Mille, Ethiopia. Proceedings of the National Academy of Sciences of the United States of America, 107 (27), 12121-6 PMID: 20566837

Kibii, J., Churchill, S., Schmid, P., Carlson, K., Reed, N., de Ruiter, D., & Berger, L. (2011). A Partial Pelvis of Australopithecus sediba Science, 333 (6048), 1407-1411 DOI: 10.1126/science.1202521

Ruff, C. (2010). Body size and body shape in early hominins – implications of the Gona Pelvis Journal of Human Evolution, 58 (2), 166-178 DOI: 10.1016/j.jhevol.2009.10.003

Simpson, S., Quade, J., Levin, N., Butler, R., Dupont-Nivet, G., Everett, M., & Semaw, S. (2008). A Female Homo erectus Pelvis from Gona, Ethiopia Science, 322 (5904), 1089-1092 DOI: 10.1126/science.1163592

Zipfel, B., DeSilva, J., Kidd, R., Carlson, K., Churchill, S., & Berger, L. (2011). The Foot and Ankle of Australopithecus sediba Science, 333 (6048), 1417-1420 DOI: 10.1126/science.1202703

Ardipithecus ramidus: mystery hips and missing legs

October 13, 2009April 18, 2024 / zcofran / 2 Comments

Let’s talk about the Ardipithecus ramidus pelvis from the partial skeleton ARA-VP-6/500. Variously preserved are a left ilium, a small part of the right ilium, and the caudal (bottom) portions of a sacrum. The fossil on which most of the reconstructions are based, the left ilium, is quite distorted and fragmented, the cracks in the bone filled in with matrix which subsequently expanded and contracted over time. The authors used CT-scans of the fossils to virtually remove adherent matrix, readjust bone to its (presumably) accurate position, and fill in cracks. The result:

(Lovejoy et al. 2009, fig. 1, p. 71e2)

Quite an odd mix of hominin and ape features, some of which are clear prior to reconstruction, others only after the reconstruction. The main things to note are the antero-lateral flare of the iliac blades, and the fairly wide and short sacrum (not clear from the photos; the sacrum is almost entirely imaginary), which are features also seen in bipedal hominins. Clear from the pictures, however, is the anterior inferior iliac spine (see bottom left inset), which is pronounced in bipeds like us, but weak/absent in apes. Finally, a bit more ape-like, is the relatively tall ischial and pubic region.

Listing individual features is all well and good if you’re into cladistics, but more interesting is the functional interpretation of the fossil and reconstruction. Here’s what the authors have to say about the ilium:

[The] exceptionally derived ilium is striking. It implies an early adaptation to habitual terrestrial bepedality before any increase in the lumbar entrapment seen in the African apes. (Lovejoy et al. 2009, p. 71e3).

Hold the phone! Why are the lateral flare and low height of the ilium necessarily adaptations to terrestrial bipedalism? One key word that follows from their reconstructions, but is not actually manifest in any of the preserved fossils, is lordosis. Lumbar lordosis refers to the frontward concavity of the lower spine, seen only in humans, fossil hominins, and bipedally-trained macaques (it’s also why we’re prone to lower back inuries).

Lordosis in Ardipithecus is entirely inferred. The (reconstructed, probably realistically) broad sacrum and shortened superior iliac blades suggest that Ardipithecus was capable of lumbar lordosis, because the lower lumbar vertebrae were not closely flanked by the adjacent ilia, as in apes. But to the best of my knowledge, they don’t mean there was lordosis. Nevertheless, Ardi’s lordosis is constantly referred to in the paper. The final word on the pelvis is that the ancestral condition of hominins (like the skull and other features, claimed to be manifest in the Ardipithecus ramidus fossils) “involved situationally dependent lordosis (during terrestrial upright walking)” (Lovejoy et al. 2009, p. 71e3). I’m not sure what exactly they mean by this, because animals can’t just adjust the relative front and back heights of their vertebrae willy-nilly. If that were the case I’d have a straight spinal column when I sleep, and overly-lordotic when I’m pregnant (which is often; Whitcome et al. 2008), and random on Halloween.

It is an interesting pelvis, though I wouldn’t be as cavalier about asserting that it belonged to an adroit terrestrial biped. Unfortunately, the partial proximal femur described with the pelvis mostly lacks any diagnostic morphology. It would be great to see what the thickness of the cortical bone in the femoral neck was like in Ardi, or the extent of articular surface on the femoral head, because these have been shown to have characteristic forms in bipeds. Hopefully future fossil discoveries will shed light on these in Ardipithecus, as well as pelvic morphology in the earliest Australopithecus.

Until then, I’m content to conclude that Ardi had a unique form of locomotion (arboreal bipedalism?), but I’m hesitant to call it a terrestrial biped.

References
Lovejoy CO, Suwa G, Spurlock L, Asfaw B, and White TD. 2009. The Pelvis and Femur of Ardipithecus ramidus: The Emergence of Upright Walking. Science 326: 71e1-71e6.

Whitcome KK, Shapiro L, and Lieberman DE. 2007. Fetal load and the evolution of lumbar lordosis in bipedal hominins. Nature 450: 1075-1080.

Tabūn Pelvis Reconstruction

April 20, 2009 / zcofran / 1 Comment

News: Weaver and Hublin (2009) virtually reconstructed the Tabūn C1 female Neandertal pelvis using CT scans.

Background: This is the closest we have to a complete female Neandertal pelvis, so a lot of the discussion centers around obstetrics. When a modern human woman gives birth, the infant enters the birth canal facing sideways so that the head will fit through the transversely oval inlet, then turns 90 degrees so it is facing the back so that the head will fit through the AP oval midplane and outlet, and finally turns another 90 degrees after the head passes through the outlet so that the shoulders can also fit through the outlet.

Tabūn conclusions: Neandertal infants (based on Tabūn’s inlet, midplane, and outlet diameters) only required two rotations: the initial turn so the head faces laterally, and the last turn so the shoulders fit through the transversely oval outlet. This means the infant comes out with the head facing sideways and the shoulders facing front (this is also how australopithecines are thought to give birth). This, the authors suggest, means that Neandertals were more primitive than modern humans. Furthermore, the transversely oval birth canal reflects the cold-adapted wide pelvis associated with Neandertals.

Problems:

Methods: There is no sacrum for Tabun. None. It is possible to predict sacral width and thus reconstruct the inlet, but it is improbable for the outlet to be reconstructed accurately without knowing the sacrum’s length, curvature, and orientation.
Sexual dimorphism: They confuse this throughout the paper. First, they female-ize Kebara (a complete male Neandertal pelvis) by assuming that Neandertals were as sexually dimorphic as modern humans. This has been shown to be wrong previously, so it was a dumb assumption. They also find that this is not the case, making me wonder why they bothered with it in the first place. Then, they claim that the difference between Neandertals and modern humans is that Neandertals are like modern males (who have short pubic rami) when really they’re like modern females (who have long pubic rami). See Rosenberg (2007) for more discussion of this.
Cold- and warm-adaptations: They say that Neandertals were cold-adapted because of the wide birth canal, in contrast to warm-adapted modern humans from Africa. First, wide birth canals do not go hand-in-hand with wide pelves. Second, Tabūn lived in the Levant and thus did not need to be cold-adapted. Third, the Busidima female Homo erectus pelvis from Gona is also wide and also not cold-adapted. Fourth, modern humans in Africa evolved a narrow pelvis to be better adapted to the warm environment is based on… uh… KNM-WT 15000? No, wait, that’s also a H erectus, and Gona has already shown that they have wide pelves despite their climate. But what else is there to support this long-held idea? Answer: Not much.

Citations:

Weaver, TD and JJ Hublin (2009) Neandertal birth canal shape and the evolution of human childbirth. PNAS Early Edition: 1-6.
Rosenberg, KR (2007) Neandertal Pelvic Remains From Krapina: Peculiar or Primitive? Periodicum Biologorum 109(4).

Busidima female pelvis

November 17, 2008August 4, 2013 / zcofran / 3 Comments

Yay! A female, mostly complete, Homo erectus pelvis has been found (Simpson et al 2008)! I say “yay” because female pelves are the best way to learn about the effects of birthing on pelvic morphology, despite the numerous studies that claim to glean this knowledge from male pelves (I’m basing this mainly on Neandertal examples as I am admittedly not as well versed with H erectus papers).

Quick summary of the facts: Found in Gona, Ethiopia, BSN49/P27a-d (a.k.a. the Busidima pelvis) consists of both os coxae (hipbones) and the upper part of the sacrum of what has been identified as an adult female Homo erectus dated to 0.9 to 1.4 Ma. The hipbones are mostly complete, except for portions of the ilium missing on both sides and portions of the ischial tuberosities. Overall, the pelvis has been reconstructed so that measurements can be taken (see image below, taken from the Simpson et al 2008 article).

Why I’m excited: Since this fossil is female and has the first complete early Pleistocene pubis, and thus the first complete pelvic inlet, it means legitimate inferences about birthing can be made. The initial paper addresses this by exploring neonate (i.e. infant at birth) head sizes compared to inlet dimensions. The authors found that Busidima would have been capable of birthing infants 30% larger than predicted based on the Homo erectus pelvis KNM-WT 15000 (male subadult). Already this find is showing why making assumptions about birth based on male specimens is flawed! Yay!!

Why I’m concerned: The authors draw conclusions about the width of the trunk based on the bi-iliac breadth. As the picture shows, the ilia are largely reconstructed, making any measurement between the widest points on the iliac crest dubious. I’m not saying it’s a bad reconstruction; in fact it looks reasonable to me. I just believe caution must be used when drawing conclusions based on reconstructed measurements. Especially since I cannot find details on how the ilia were reconstructed (if I’ve missed something in my reading of the online supporting material, please, someone correct me). Given that this is the case, it seems extreme to use incomplete ilia as evidence against an endurance running hypothesis and as evidence for what type of environment this specimen lived in.

What others are saying: Hawks goes into more anatomical detail in his blog post about the article. He discusses questions raised by the study and points out that you really have to read the supporting stuff to get the full picture as the Science article is “superficial”. Beast Ape’s blog post discusses the bi-iliac interpretation and what it means. For a multilingual look, Mundo Neandertal also discusses the article, and based on my imperfect Spanish capabilities, I think they discuss the infant head size findings. Finally, New Scientist describes the findings in a newsy way, detailing the bigger birth canal (compared to WT 15000) and the squat proportions of Busidima. Enjoy!

References:
Simpson et al. 2008 A Female Homo erectus Pelvis from Gona, Ethiopia. Science 322: 1089-1092.

Lawn Chair Anthropology

Biological anthropology, paleontology, evolution, and development, by Dr. Zachary Cofran

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