Today we’re delighted to have a guest post from Dr. Chris Noto, a new assistant professor at University of Wisconsin-Parkside, and an old friend of mine from our graduate school days together at Stony Brook University. Chris has had a long-running interest in dinosaur paleoecology, and thus it only seemed natural for him to apply these interests to the ODP data. Enjoy!
In describing the ecology of an organism our first inclination may be to simply go observe it in its day-to-day existence. Therefore, at its core, ecology is primarily a science of the living and this is reflected in the methods and theories one finds in the literature. Over the past couple decades there has been a growing interest in the relationship between organismal morphology and ecology, which is now often referred to as “ecomorphology”(Losos 1990; Ben-Moshe et al. 2001; Aguirre et al. 2002; Zeffer et al. 2003; Sacco and Van Valkenburgh 2004). This has opened entire new areas of research into the covariation between organisms and their environment, which is also the basic foundation for understanding evolutionary change over longer spans of time.
Some paleontologists have applied ecomorphological principles to reconstructing the paleoecology of certain extinct groups, including carnivorans (Palmqvist et al. 1999), birds (Hertel 1995), and especially ungulates (Solounias and Semprebon 2002; Meehan and Martin 2003; DeGusta and Vrba 2005; Klein et al. 2010). Changes in the types and/or proportions of ecomorphs in a fossil community have also been used as evidence of environmental and evolutionary responses to climate change(Van Valkenburgh 1995; Meehan and Martin 2003; Badgley et al. 2008; Noto and Grossman 2010). You will note though that a lot of this research relies on comparison to living analogs related to the fossil groups in question. How do we explore the paleoecology of groups that completely lack extant analogs?
If there’s one thing we’ve learned through all this research, it’s the fact that:
- Certain morphological adaptations occur regardless of species (convergence) because of specific habitat constraints, and
- Morphological differences between species will occur due to diverging ecologies, even if we don’t know exactly what ecological functions those morphological differences actually represent.
But, we won’t know those differences exist until we look for them. Paleoecology is first and foremost comparative: we take our fossils and compare them to other related taxa and living forms to better understand their place in the original community. Often we assign categories to taxa, such as “carnivore”, “biped”, etc.; however, differences between species are often better described by a continuum than a set of categories(Carrano 1999). The morphology of an organism reflects the amount of time it spends doing certain activities or performing certain functions. For example, a sloth can swim on occasion even if it is not particularly well adapted for it. Dinosaur paleoecology is finally moving in the direction of our mammalian colleagues by using quantitative measures of morphology (which allow for continua) instead of assigning discreet categories.
The ODP is one of the first large-scale projects to bring together the kind of dataset necessary to study dinosaur ecomorphology. In a recent paper I published looking at differences between dinosaur fossil communities (Noto and Grossman 2010), I was forced to use categories in assigning ecomorphs, which artificially restricted the analysis by forcing me to choose a category when uncertainty existed. In this case it was whether certain non-hadrosaur ornithopods were bipedal or quadrupedal. With ODP data, it is now possible to take a more quantitative (and nuanced) approach to this question.
To explore possible trends in ornithischians, I used humerus length and mediolateral width measurements to calculate Mike Taylor’s Gracility Index (GI; Taylor 2009) using only the largest individual from each species. These data were log transformed and plotted against the log of humerus length (to help minimize the effects of size and codependency). The resulting plot clearly separates the taxa, with more bipedal taxa having relatively gracile humeri and quadrupedal groups have more robust humeri. There are two ways to use this graph. First, we can look at the distribution of taxa from each group and see whether they fit more towards a bipedal or quadrupedal type; those intermediate to the extremes are referred to as facultative. These are my own divisions based on where I see breaks in the data. Basal ceratopsians, for example, occur mainly towards the bipedal or facultative ends of the spectrum, while derived Neoceratopsians are firmly on the quadrupedal end of things. Another way to look at the plot is how robust we may expect the humerus to be for a taxon of a given size. The dashed line is drawn across the middle of the plot. For a given humerus length we can compare GI between taxa. For example, Iguanodon has a more robust humerus (lower GI) than most ornithopods in the dataset. Furthermore, we can spot outliers, which may point to either extreme specialization or faulty data. The theropod Mononykus has an extremely robust humerus, approaching the level of Triceratops, which is related to its specialized digging forelimb. On the other hand, Cerasinops appears to have the most robust humerus of all, however as Mike pointed out to me, the original paper describes the humerus as extremely gracile and gives no width measurement. So where did the width measurement come from? This particular data point is worth another look.
As you can see, the distribution of humeral morphologies indicates a gradual continuum of locomotor strategies from fully bipedal to full quadrupedal. Quantitative data such as this could then be fed into a paleoecological analysis instead of categories, allowing for more refined analysis of ecological differences and similarities among paleocommunities over space and time. While evolutionary trends are certainly important, we must not forget the ecological context of the morphological patterns we are studying.
Aguirre LF, Herrel A, van Damme R et al. (2002) Ecomorphological analysis of trophic niche partitioning in a tropical savannah bat community. Proceedings of the Royal Society of London Series B-Biological Sciences 269:1271-1278
Badgley C, Barry JC, Morgan ME et al. (2008) Ecological changes in Miocene mammalian record show impact of prolonged climatic forcing. Proc Natl Acad Sci U S A 105:12145-12149
Ben-Moshe A, Dayan T, Simberloff D (2001) Convergence in morphological patterns and community organization between Old and New World rodent guilds. Am Nat 158:484-495
Carrano MT (1999) What, if anything, is a cursor? Categories versus continua for determining locomotor habit in mammals and dinosaurs. J Zool 247:29-42
DeGusta D, Vrba E (2005) Methods for inferring paleohabitats from discrete traits of the bovid postcranial skeleton. J Archaeol Sci 32:1115-1123
Hertel F (1995) Ecomorphological indicators of feeding behavior in recent and fossil raptors. Auk 112:890-903
Klein RG, Franciscus RG, Steele TE (2010) Morphometric identification of bovid metapodials to genus and implications for taxon-free habitat reconstruction. J Archaeol Sci 37:389-401
Losos JB (1990) Ecomorphology, performance capability, and scaling of West Indian Anolis lizards: an evolutionary analysis. Ecol Monogr 60:369-388
Meehan TJ, Martin LD (2003) Extinction and re-evolution of similar adaptive types (ecomorphs) in Cenozoic North American ungulates and carnivores reflect van der Hammen’s cycles. Naturwissenschaften 90:131-135
Noto CR, Grossman A (2010) Broad-scale patterns of Late Jurassic dinosaur paleoecology. PLoS ONE 5:e12553
Palmqvist P, Arribas A, Martinez-Navarro B (1999) Ecomorphological study of large canids from the lower Pleistocene of southeastern Spain. Lethaia 32:75-88
Sacco T, Van Valkenburgh B (2004) Ecomorphological indicators of feeding behaviour in the bears (Carnivora : Ursidae). J Zool 263:41-54
Solounias N, Semprebon G (2002) Advances in the reconstruction of ungulate ecomorphology with application to early fossil equids. Am Mus Novit:1-49
Taylor M (2009) A re-evaluation of Brachiosaurus altithorax Riggs 1903 (Dinosauria, Sauropoda) and its generic separation from Giraffatitan brancai (Janensch 1914). J Vert Paleontol 29:787-806
Van Valkenburgh B (1995) Tracking ecology over geological time – evolution within guilds of vertebrates. Trends Ecol Evol 10:71-76
Zeffer A, Johansson LC, Marmebro A (2003) Functional correlation between habitat use and leg morphology in birds (Aves). Biol J Linn Soc 79:461-484
Local disparity guru and paleontologist Randy Irmis (that’s Randall B. Irmis, Ph.D., if you go by his web page) recently posted a nice long list of recommended readings on the issue of disparity – what it is, how to calculate it, etc. As a reminder, disparity is the measure of how different species are from each other in terms of shape, size, or other discrete features (not the same as diversity, which just counts how many different species exist – once again, see Randy’s eloquent post on the topics). It just so happens that documenting disparity in ornithischian dinosaurs is at the top of our list for the ODP. Hence, I decided to buckle down and read through an important recent paper on the topic (one that Randy happened to highlight in his list, too).
In the interest of getting this post out in a timely manner, I’m mainly going to be posting my unpolished notes, taken a few weeks ago in the comfort of my bed (nothing like a little light bedtime reading). I’ve made a few adjustments here and there, but otherwise you can consider this a peek into my stream of consciousness while reading the literature. Because I was mainly interested in how the work could be applied to the ODP, I didn’t really bother with summarizing the specific analyses done by the authors. Thus, without further preface:
Brusatte, S. L., Montanari, S., Yi, H.-Y, and Norell, M. A. 2011. Phylogenetic corrections for morphological disparity analysis: new methodology and case studies. Paleobiology 37: 1-22. [unfortunately, not openly available as a PDF] [link to abstract]
The Main Gist:
The fossil record just isn’t complete – and that’s particularly true for many of the early members of important ornithischian clades (like thyreophorans and marginocephalians). However, it’d be nice to interpolate some of these missing data in order to produce a more complete picture of the changes in a clade’s disparity over time and in morphospace (the multi-dimensional plot of the shape of an animal’s bones, in this case). Brusatte and colleagues, building on the work of many other authors, have formalized a method to fill in some of these gaps by producing a plausible reconstruction of missing ancestors.
[as presented here, it's a mix of to-do tasks for the ODP, a cookbook for the analysis, and how Brusatte et al.'s method will be applied; caveat emptor]
The questions: What is the morphospace occupied by ornithischian dinosaurs over time? How does the morphospace change? How does the morphospace occupied by specific clades differ?
- Assemble data matrix (taxon/measurement matrix)
- Reconstruct ancestoral measurements following Brusatte et al. 2011
- Calculate Euclidean distance matrix (“quantifies the pairwise dissimilarity between taxa”) – this presumably calculates dissimilarity for each taxon/measurement pair
- Apply principal coordinates analysis (PCoA) to each analysis (better handles missing data than does PCA [principal components analysis]). Can be done in R.
- PCoA produces scores for each taxon along n=#taxa axes. Can be done in R.
- Examine slope of scree plot to determine where break occurs; only examine these “interesting” axes. I think this scree plot can be done in R.
- Calculate disparity indices from the PCoAs, using different bins (categories). Can be done in R. Categories might include: 1) clade; 2) time; 3) locomotor category; 4) combination of clade/locomotor category.
- Indices include: sum of range of values along axis 1, 2, … n (i.e., range 1+range2+range3. . .); product of range of values along axis 1, 2, n (range 1 * range 2 * range 3. . .) normalized to the nth root; and same sum and products for variance in each bin.
- Rinse and repeat using ancestral values as calculated following Brusatte et al. 2011.
- Disparity indices that can be compared statistically (using bootstrap values) for various categories. E.g., a disparity value for Ceratopsia, Ornithopoda, Thyreophora, etc. disparity value for quadrupeds vs. bipeds.
- Graphs showing point clouds for various clades along various axes (e.g., PC1 vs. PC2)
- Graphs showing trends for disparity over time, with different groups. E.g., trend line showing disparity in ornithischians as a whole, along with trend line showing disparity in thyreophorans, ceratopsians, etc. Potential sample size issues here, particularly for clades with few members or few members early in their history
- Narrative text and / or table showing what factors are loaded on which axes
In response to a recent query on this blog, ODPer Christian Foth contributed a list of papers potentially relevant to the ODP, specifically limb posture and evolution in ornithischian dinosaurs. It’s important to recognize work that others did before and see how it relates to ours. Furthermore, a good reference list is essential for the upcoming manuscript.
What can you do?
If you think of another paper that might be added to the list (within reason, of course), drop a line in the comments section. If you are interested in providing a summary of a certain paper as a guest blog post (either here or at your own blog), that would be great, too. As always, one need not be a Ph.D’ed scientist to apply! We’re just looking for a short summary.
For my part, I added the Middleton and Gatesy reference – although it deals with theropods, I think some of the background info and analytical methods are quite relevant. Hmm. . .that might be a good one to blog about.
Reference List in Progress
Alexander R. 1985. Mechanics of posture and gait of some large dinosaurs. Zoological Journal of the Linnean Society 83: 1-25.
Bakker RT. 1968. The superiority of dinosaurs. Discovery 3 (2): 11-22.
Biewener (1983). Allometry of quadrupedal locomotion: the scaling of duty factor, bone curvature and limb orientation to body size. J. Exp. Biol. 105: 147-171.
Bonnan MF & P Senter. 2007. Were the basal sauropodomorph dinosaurs Plateosaurus and Massospondylus habitual quadropeds. Special Papers in Palaeontology 77: 139–155
Bonnan, MF, & AM Yates. 2007. A new description of the forelimb of the basal sauropodomorph Melanorosaurus: implications for the evolution of pronation, manus shape and quadrupedalism in sauropod dinosaurs. pp. 157-168 in: Paul M. Barrett and David J. Batten (eds.), Special Papers in Palaeontology 77: Evolution and Palaeobiology of Early Sauropodomorph Dinosaurs. The Palaeontological Association, U.K.
Bultynck P (1992) An assessment of posture and gait in Iguanodon bernissartensis Boulenger, 1881. Bulletin de l’Institut Royal des Sciences Naturelles de Belgique: Sciences de la Terre 63: 5-11.
Carrano MT. 2000. Homoplasy and the evolution of dinosaur locomotion. Paleobiology 26 (3): 489-512.
Carrano MT (2001) Implications of limb bone scaling, curvature and eccentricity in mammals and non-avian dinosaurs. Journal of Zoology 254: 41-55.
Dilkes DW. 2000. Appendicular myology of the hadrosaurian dinosaur Maiasaura peeblesorum from the Late Cretaceous (Campanian) of Montana. Transactions of the Royal Society of Edinburgh, Earth Sciences 90: 87-125.
Dilkes DW. 2001. An ontogenetic perspective on locomotion in the Late Cretaceous dinosaur Maiasaura peeblesorum (Ornithischia: Hadrosauridae). Canadian Journal of Earth Sciences 38: 1205-1227.
Dodson, P & JO Farlow. 1997. The forelimb carriage of ceratopsid dinosaurs. DinoFest International Proceedings 393-398.
Galton PM. 1970. The posture of hadrosaurian dinosaurs. Journal of Paleontology 44 (3): 464-473.
Garstka WR & DA Burnham. 1997. Posture and stance of Triceratops. Evidence of digitigrade manus and cantilever vertebral column. DinoFest International Proceedings 385-391.
Heinrich DE, Bruff CB & DB Weishampel. 1993. Femoral ontogeny and locomotor biomechanics of Dryosaurus lettowvorbecki (Dinosauria, Iguanodontia). Zoological Journal of the Linnean Society 108: 179-196.
Hutchinson JR. 2004. Biomechanical modeling and sensitivity analysis of bipedal running ability. I. Extant taxa. Journal of Morphology 262: 421-440.
Johnson RE, Ostrom JH (1995) The forelimb of Torosaurus and an analysis of the posture and gait of ceratopsian dinosaurs. In: Thomason JJ, editor. Functional Morphology in Vertebrate Paleontology. New York: Cambridge University Press. pp. 205-218.
Kilbourne BM & PJ Makovicky. 2010. Limb bone allometry during postnatal ontogeny in non-avian dinosaurs. Journal of Anatomy 217: 135-152.
Kubo T & MJ Benton. 2007. Evolution of hindlimb posture in archosaurs: limb streeses in extinct vertebrates. Palaeontology 50 (6): 1519-1529.
Lee DV & SG Meek 2005. Directionally compliant legs influence the intrinsic pitch behaviour of a trotting quadroped. Proceedings of the Royal Society B 272(1563): 567–572.
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McMahon, T (1975) Allometry and biomechanics: Limb bones in adult Ungulates. Am. Nat. 109:547-563.
Middleton KM & S Gatesy. 2000. Theropod forelimb design and evolution. Z J Linn Soc 128: 149-187
Organ CL. 2006. Biomechnics of ossified tendons in ornithopod dinosaurs. Paleobiology 31 (4): 652-665.
Papantoniou V, Avlakiotis P & R Alexander. 1999. Control of a robit dinosaur. Phil.Trans. R. Soc. Lond. B 354: 863-868.
Paul GS & P Christiansen. 2000. Forelimb posture in neoceratopsian dinosaurs: implications for gait and locomotion. Paleobiology, 26 (3): 450–465.
Romer AS. 1923. The ilium in dinosaurs and birds. Bulletin American Museum of Natural Histroy 48: 141-145.
Raichlen DA. 2006. Effects of limb mass distribution on mechanical power outputs during quadrupedalism. The Journal of Experimental Biology 209: 633-644
Romer AS. 1927. The pelvic musculature of ornithischian dinosaurs. Acta Zoologica 8: 225-275.
Sellers WI & PL Manning. 2007. Estimating dinosaur maximum running speeds using evolutionary robotics. Proceedings of Royal Society B 274: 2711-2716.
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Sternberg CM. 1965. New restoration of hadrosaurian dinosaur. National Museum of Canada, Natural History Papers 1-5.
Taylor CR. 1978. Why change gaits? Recruitment of muscles and muscle fibers as a function of speed and gait. American Zoologist 18: 153.161.
Tereshchenko VS. 1994. A reconstruction of the erect posture of Protoceratops. Paleontological Journal 28 (1): 104-119.
Tereshchenko VS. 1996. A reconstruction of the locomotion of Protoceratops. Paleontological Journal 30 (2): 232-245.
Tereshchenko VS. 2008. Adaptive features of protoceratopoids (Ornithischia: Neoceratposia). Paleontological Journal 42 (3): 273-286.
Thompson S & R Holmes. Forelimb stance and step cycle in Chasmosaurus irvenenesis (Dinosauria: Neoceratopsia). Palaeontologica Electronica 10 (1): 5A.
Thulborn RA. 1982. Speeds and gaits of dinosaurs. Palaeogeography, Palaeoclimatology, Palaeoecology, 38: 227-256.
Thulborn RA. 1984. Preferred gaits of bipedal dinosaurs. Alcheringa 8 (3): 243-252.
Yates, Adam M., Matthew F. Bonnan, Johann Neveling, A. Chinsamy and Marc G. Blackbeard. 2009. A new transitional sauropodomorph dinosaur from the Early Jurassic of South Africa and the evolution of sauropod feeding and quadrupedalism. Proceedings of the Royal Society B, published online. doi:10.1098/rspb.2009.1440
Dedicated readers of the blog likely remember that one of the core research goals of this project is to examine the bipedal/quadrupedal transition in ornithischian dinosaurs. Of course, ornithischians weren’t the only group to experience such locomotor changes during their evolution! A new paper on the bipedal/quadrupedal transition in sauropodomorphs (the saurischian dinosaur group including animals like Plateosaurus, Apatosaurus, and Brachiosaurus) has just appeared on Proceedings of the Royal Society B‘s FirstCite. This paper, headed up by Adam Yates, details the anatomy of Aardonyx, an early sauropodomorph from the Early Jurassic of South Africa. The authors posit that the new critter was a habitual biped, although it had many features that presaged the anatomy of later quadrupedal forms.
If you haven’t contributed data to the Open Dinosaur Project yet, and are looking for something to do, this might be a good one! The supplementary information (freely available) is chock-full of measurements.
Yates, A. M., Bonnan, M. F., Neveling, J., Chinsamy, A., and M. G. Blackbeard. In press. A new transitional sauropodomorph dinosaur from the Early Jurassic of South Africa and the evolution of sauropod feeding and quadrupedalism. Proceedings of the Royal Society B. doi:10.1098/rspb.2009.1440. Published online 10 November 2009. [subscription required for full access]