More Tininess From the Malay Archipelago

I've been blogging on nothing but code lately, so here's a "quick" paleo-post.

"Hobbits": Primitives or Pinheads?

The anthropological world was shocked several years ago by the discovery of a relatively recent pygmy species of stem-human, popularly dubbed "hobbits" and more technically dubbed Homo floresiensis (Brown et al. 2004). Known from subfossil remains of several individuals from the island of Flores (Morwood, Brown et al. 2004), including one nearly complete skeleton including the skull (the holotype specimen), these people stood barely over a meter in height, shorter than any modern pygmy humans. They survived at least until 13,000 years before present (Morwood, Soejono et al. 2004), and, judging from the one skull, they had extremely small brains for such late-surviving Homo (380 cc—we average around 1400 cc and even Australopithecus africanus averaged in the 400s). They had other primitive features as well, such as chinless jaws, a low twist in the forearm bones (Ibid.), robust leg bones (Ibid.), a chimp-like wrist (Tocheri et al. 2007), and some features of the teeth.

But not everyone agrees that the Floresian "hobbits" are a separate species. Various researchers have proposed that they are, in fact, pygmy Homo sapiens and that at least some individuals, including the holotype, have a pathological condition that includes microcephaly, a.k.a. "pinheadedness" (Martin et al. 2006, Jacob et al. 2006). After all, at least some of the supposedly diagnostic features, such as chinlessness, are present in some members of our own species (Jacob et al. 2006). But others insist that there are genuine primitive features in the Floresian specimens that cannot be explained by microcephaly or dwarfism (Falk et al. 2005, Falk et al. 2006, Argue et al. 2006).

A New Discovery

Until recently, the Floresians' size distinguished them from H. sapiens. But that was before this week's publication (Berger et al. 2008; open access) of some very recent (1400 to 2900 years B.P.) pygmy/dwarf fossils from Palau. More work needs to be done to free these fossils from the limestone they were found in, but they seem to be Homo sapiens proper, high-domed skulls and all. What's really interesting is that, like the Floresian "hobbits", the Palauans are smaller than any living population of H. sapiens. But unlike the "hobbit" holotype, their cranial capacity is not abnormally low.

Berger et al. note that the Palau specimens have some supposedly "primitive" features that may in fact be related to size, such as proportionally large teeth (megadontia) and chinless mandibles. The authors don't firmly ally themselves with either camp, but note that at least some of the supposed diagnostic features of H. floresiensis are probably phylogenetically uninformative, merely being results of dwarfism.

Thoughts

Personally, while I'm no expert in any the fields involved, I find the arguments that the Floresians are pathological, pygmy H. sapiens unconvincing. For one thing, people afflicted with the proposed diseases don't actually look like the "hobbits" except in having small brains. For another, asymmetry that was ascribed to pathology seems more likely to me to be a preservational artifact. Look at just about any fossil skull face-on—most of them are asymmetrical.

But more damning are the limb characters, which no known pathology can explain. (Did they have some kind of "ape-itis" in addition to microcephaly?) Fact is, I think that if H. floresiensis were 130,000 years old instead of 13,000, nobody would even contest the idea that they weren't H. sapiens. And morphology deserves more weight than chronology in matters of systematics.

References

• Argue, D., D. Donlon, C. Groves and R. Wright (2006 October 4). Homo floresiensis: microcephalic, pygmoid, Australopithecus, or Homo? Journal of Human Evolution 51(4):360–374. doi:10.1016/j.jhevol.2006.04.013
• Berger, L. R., S. E. Churchill, B. De Klerk and R. L. Quinn (2008 March 12). Small-bodied humans from Palau, Micronesia. PLoS ONE 3(3):e1780. doi:10.1371/journal.pone.0001780
• Brown, P., T. Sutkina, M. J. Morwood, R. P. Soejono, Jatmiko, E. Wayhu Saptomo and Rokus Awe Due. (2004 October 28). A new small-bodied hominin from the Late Pleistocene of Flores, Indonesia. Nature 431:1055–1061. doi:10.1038/nature02999
• Falk, D., C. Hildebolt, K. Smith, M. J. Morwood, T. Sutkina, P. Brown, Jatmiko, E. Wayhu Saptomo, B. Brunsden and F. Prior (2005 April 8). The brain of LB1, Homo floresiensis. Science 308(5719):242–245. doi:10.1126/science.1109727
• Falk, D., C. Hildebolt, K. Smith, M. J. Morwood, T. Sutkina, P. Brown, Jatmiko, E. Wayhu Saptomo, B. Brunsden and F. Prior (2006 May 19). Response to comment on "The brain of LB1, Homo floresiensis". Science 312(5776):999. doi:10.1126/science.1124972
• Jacob, T. E., Indriati, R. P. Soejono, K. Hsü, D. W. Frayer, R. B. Eckhardt, A. J. Kuperavage, A. Thorne and M. Hennenberg (2006 September 5). Pygmoid Australomelanesian Homo sapiens skeletal remains from Liang Bua, Flores: population affinities and pathological abnormalities. Proceedings of the National Academy of Sciences of the United States of America 103(36):13421–13426. doi:10.1073/pnas.0605563103
• Martin, R. D., A. M. MacLarnon, J. L. Philips, L. Dussubieux, P. R. Williams and W. B. Dobyns (2006 May 19). Comment on "The brain of LB1, Homo floresiensis". Science 312(5776):999. doi:10.1126/science.1121144
• Morwood, M. J., P. Brown, Jatmiko, T. Sutkina, E. Wahyu Saptomo, K. E. Westaway, Rokus Awe Due, R. G. Roberts, T. Maeda, S. Wasisto and T. Djubiantono (2005 October 13). Further evidence for small-bodied hominins from the Late Plesitocene of Flores, Indonesia. Nature 437:1012–1017. doi:10.1038/nature04022
• Morwood, M. J., R. P. Soejono, R. G. Roberts, T. Sutkina, C. S. M. Turney, K. E. Westaway, W. J. Rink, J.-x. Zhao, G. D. van den Bergh, Rokus Awe Due, D. R. Hobbs, M. W. Moore, M. I. Bird and L. K. Fifield (2004 October 28). Archaeology and age of a new hominin from Flores in eastern Indonesia. Nature 431:1087–1091. doi:10.1038/nature02956
• Tocheri, M. W., C. M. Orr, S. G. Larson, T. Sutikna, Jatmiko, E. Wahyu Saptomo, Rokus Awe Due, T. Djubiantono, M. J. Morwood and W. L. Jungers (2007 September 21). The primitive wrist of Homo floresiensis and its implications for hominin evolution. Science 317(5845):1743–1745. doi:10.1126/science.1147143

Names on NEXUS: Under the Hood

I nearly have the basic data model and data processing functions pinned down for Names on NEXUS. Once again, that's my project, hinted at in a paper of mine (Keesey 2007), to relate the data in NEXUS files (Maddison et al. 1997) to definitions of names as governed by the PhyloCode.

I've had to learn some new technologies and code packages to accomplish this. Here's a rundown of some key ones:

BioJava
This is the most recent addition. Originally I had built my own library in ActionScript 3.0 to parse NEXUS files. But it had some limitations. NEXUS is a rather old format (as bioinformatics formats go), and different applications produce somewhat different versions. So rather than use my own ad hoc library, I decided I should get an open source one.

There aren't any in ActionScript, of course, but there are some in Java. This meant I had to switch NEXUS parsing from the front end to the back end, but in some ways that's better. It means I can stored parsed data in the database instead of having client application parse NEXUS data every time. In fact, it means that the client never has to actually see raw NEXUS data—it can just fetch the pre-parsed data.

I first looked into using the NEXUS-parsing code in Mesquite, an open-source phylogenetic analysis program. But it's not set up for simply using the parsing engine on its own—the parser is tied into a whole file-browsing package. Then I found BioJava, which had exactly what I needed. Just looka this package!

Unfortunately there are still some problems with opening certain NEXUS files. I downloaded some samples from TreeBASE and they flagged errors in the TREES section. The reason, as I found after hours of searching and considering whether it might be better just to write my own parser after all, turns out to be an extra comma in the TRANSLATE section. Still not exactly sure how I'm going to solve that one. But it works when I remove the comma!

Hibernate
Remember how I wrote a post a while ago about building classes that map from the Java back-end to the database? Turns out that was all unnecessary. Hibernate is a persistence layer that provides pretty seamless integration between Java and a database (in this case, a PostgreSQL database). Augmented by Hibernate Annotations and Hibernate Validator, it makes it fairly easy to set up and use a complex, well-organized database.

Well, okay, there's a bit of a learning curve first, but it's totally worth it. Incidentally, the book I used to learn it has what is possibly the best title ever.

Flex Data Management Services
Basically, Hibernate is to Java and databases as mx.data is to Flex and Java. It provides a persistence layer so that I don't have to keep track of whether or not I need to request certain data from the Java back-end. I just create DataService objects, tie them to Assembler classes on the back end, and it's all taken care of.

FlexUnit and JUnit
I've already extolled the virtues of unit testing. These wonderful (and, yes, comically-named) packages (huh huh) make it possible. I haven't built enough unit tests, really, but the few I have have been enormously useful in hunting down peculiar errors. And aside from that, since Eclipse can run JUnit tests natively, I can even use them to perform certain important tasks, such as setting up the database from annotated classes via Hibernate.

So What's Left For Me To Do?
Plenty. Although these premade packages help out enormously, I've still had to build an entire mathematics library, a MathML parser, and some tools for handling URIs. I've still got tons of work left to do on the user interface. (Event bubbling is helping a lot with that, by the way.) And, even when stuff is already built, just hooking up one pipe to another pipe can be more complicated than it seems.

Here's a rough list of what's left:

• Finalize the servlet for uploading and parsing NEXUS data. (I'm very close on this one.)
• Finish the required behind-the-scenes "search" features. Some of these might be a bit involved, like the ones that suggest possible links between NEXUS taxa and species or specimens or between NEXUS character states and apomorphies.
• Overhaul the way Names on NEXUS entities (particularly specifiers) are referenced in MathML.
• Finish the user interface. So far I just have a few forms. I still have to do tree visualization, stylesheets, high-level navigation, transitions, etc.
• Constrain access to certain functionality. Names on NEXUS is going to be a pretty open, collaborative tool, but I need to set a few boundaries. (E.g., I can't have any old person delete data.)
• Make sure the server's all optimized, with a static, JNDI-named Hibernate factory, etc.

And here are some things that aren't, strictly speaking, essential, but would be awfully nice:

• Create a servlet to provide permanent links for Names on NEXUS entities.
• Create unit tests for all relevant classes.
• Add JavaDoc and ASDoc comments to all code.

Part of me is also thinking about renaming the project. I mean, it's a good name for what it does right now, but what if I start to bring formats other than NEXUS into the fold? (Not that there are many, but....) Well, I'll probably cross that bridge when I come to it.

My goal is to get an alpha version online sometime this Spring and go open source with it by the Fall. We'll see....