First anatomical network analysis of fore- and hindlimb musculoskeletal modularity in bonobos, common chimpanzees, and humans

Studies of morphological integration and modularity, and of anatomical complexity in human evolution typically focus on skeletal tissues. Here we provide the first network analysis of the musculoskeletal anatomy of both the fore- and hindlimbs of the two species of chimpanzee and humans. Contra long-accepted ideas, network analysis reveals that the hindlimb displays a pattern opposite to that of the forelimb: Pan big toe is typically seen as more independently mobile, but humans are actually the ones that have a separate module exclusively related to its movements. Different fore- vs hindlimb patterns are also seen for anatomical network complexity (i.e., complexity in the arrangement of bones and muscles). For instance, the human hindlimb is as complex as that of chimpanzees but the human forelimb is less complex than in Pan. Importantly, in contrast to the analysis of morphological integration using morphometric approaches, network analyses do not support the prediction that forelimb and hindlimb are more dissimilar in species with functionally divergent limbs such as bipedal humans

A Field Approach to 3D Gene Expression Pattern Characterization

We present a vector field method for obtaining the spatial organization of 3D patterns of gene expression based on gradients and lines of force obtained by numerical integration. The convergence of these lines of force in local maxima are centers of gene expression, providing a natural and powerful framework to characterize the organization and dynamics of biological structures. We apply this novel methodology to analyze the expression pattern of the Enhanced Green Fluorescent Protein (EGFP) driven by the promoter of light chain myosin II during zebrafish heart formation.Comment: 9 pages, 2 figures, The following article has been submitted to Applied Physics Letters. If it is published, it will be found online at http://apl.aip.or

Anatomical Network Comparison of Human Upper and Lower, Newborn and Adult, and Normal and Abnormal Limbs, with Notes on Development, Pathology and Limb Serial Homology vs. Homoplasy

How do the various anatomical parts (modules) of the animal body evolve into very different integrated forms (integration) yet still function properly without decreasing the individual's survival? This long-standing question remains unanswered for multiple reasons, including lack of consensus about conceptual definitions and approaches, as well as a reasonable bias toward the study of hard tissues over soft tissues. A major difficulty concerns the non-trivial technical hurdles of addressing this problem, specifically the lack of quantitative tools to quantify and compare variation across multiple disparate anatomical parts and tissue types. In this paper we apply for the first time a powerful new quantitative tool, Anatomical Network Analysis (AnNA), to examine and compare in detail the musculoskeletal modularity and integration of normal and abnormal human upper and lower limbs. In contrast to other morphological methods, the strength of AnNA is that it allows efficient and direct empirical comparisons among body parts with even vastly different architectures (e.g. upper and lower limbs) and diverse or complex tissue composition (e.g. bones, cartilages and muscles), by quantifying the spatial organization of these parts-their topological patterns relative to each other-using tools borrowed from network theory. Our results reveal similarities between the skeletal networks of the normal newborn/adult upper limb vs. lower limb, with exception to the shoulder vs. pelvis. However, when muscles are included, the overall musculoskeletal network organization of the upper limb is strikingly different from that of the lower limb, particularly that of the more proximal structures of each limb. Importantly, the obtained data provide further evidence to be added to the vast amount of paleontological, gross anatomical, developmental, molecular and embryological data recently obtained that contradicts the long-standing dogma that the upper and lower limbs are serial homologues. In addition, the AnNA of the limbs of a trisomy 18 human fetus strongly supports Pere Alberch's ill-named "logic of monsters" hypothesis, and contradicts the commonly accepted idea that birth defects often lead to lower integration (i.e. more parcellation) of anatomical structures

Primate modularity and evolution: first anatomical network analysis of primate head and neck musculoskeletal system

Network theory is increasingly being used to study morphological modularity and integration. Anatomical network analysis (AnNA) is a framework for quantitatively characterizing the topological organization of anatomical structures and providing an operational way to compare structural integration and modularity. Here we apply AnNA for the first time to study the macroevolution of the musculoskeletal system of the head and neck in primates and their closest living relatives, paying special attention to the evolution of structures associated with facial and vocal communication. We show that well-defined left and right facial modules are plesiomorphic for primates, while anthropoids consistently have asymmetrical facial modules that include structures of both sides, a change likely related to the ability to display more complex, asymmetrical facial expressions. However, no clear trends in network organization were found regarding the evolution of structures related to speech. Remarkably, the increase in the number of head and neck muscles – and thus of musculoskeletal structures – in human evolution led to a decrease in network density and complexity in humans

Unusually thick dinosaur eggshell fragments from the Spanish Late Cretaceous

[EN] Fieldwork carried out recently in the southeastern branch of the Iberian Range (Valencia Province, Spain) has led to the collection of a large volume of dinosaur eggshell fragments of unusual thickness. These specimens, up to 4.9 mm thick, were recovered from palustrine grey marls of the upper Campanian-lower Maastrichtian Sierra Perenchiza Formation, which comprises a wetland paleoenvironment deposit. These eggshell fragments have a characteristic compactituberculate ornamentation, dinosauroid-spherulitic organisation, and exhibit a complex canaliculate respiratory system. The external tuberculate surface of the shell as well as the internal microstructure enable referral to Megaloolithus aff. siruguei, the most common megaloolithid oospecies known from the Iberian Peninsula and southern France. The biostratigraphic range of M. siruguei matches the temporal distribution of titanosaurid dinosaurs across the Iberian Range, tentatively considered to be potential producers.This work was supported by the Ministerio de Economia y Competitividad of Spain [Secretaria de Estado de Investigacion, Desarrollo e Innovacion, projects CGL2013-47521-P and CGL2014-53548-P]Company Rodríguez, J. (2017). Unusually thick dinosaur eggshell fragments from the Spanish Late Cretaceous. Historical Biology (Online). 31(2):203-210. https://doi.org/10.1080/08912963.2017.1357717S203210312Allain, R., & Suberbiola, X. P. (2003). Dinosaurs of France. Comptes Rendus Palevol, 2(1), 27-44. doi:10.1016/s1631-0683(03)00002-2Bravo, A. M., & Gaete, R. (2014). Titanosaur eggshells from the Tremp Formation (Upper Cretaceous, Southern Pyrenees, Spain). Historical Biology, 27(8), 1079-1089. doi:10.1080/08912963.2014.934231Canudo, J. I., Oms, O., Vila, B., Galobart, À., Fondevilla, V., Puértolas-Pascual, E., … Blanco, A. (2016). The upper Maastrichtian dinosaur fossil record from the southern Pyrenees and its contribution to the topic of the Cretaceous–Palaeogene mass extinction event. Cretaceous Research, 57, 540-551. doi:10.1016/j.cretres.2015.06.013Cruzado-Caballero, P., Ruiz-Omeñaca, J. I., Gaete, R., Riera, V., Oms, O., & Canudo, J. I. (2013). A new hadrosaurid dentary from the latest Maastrichtian of the Pyrenees (north Spain) and the high diversity of the duck-billed dinosaurs of the Ibero-Armorican Realm at the very end of the Cretaceous. Historical Biology, 26(5), 619-630. doi:10.1080/08912963.2013.822867Chiappe, L. M., Coria, R. A., Dingus, L., Jackson, F., Chinsamy, A., & Fox, M. (1998). Sauropod dinosaur embryos from the Late Cretaceous of Patagonia. Nature, 396(6708), 258-261. doi:10.1038/24370Company J. 2004. Vertebrados continentales del Cretácico superior (Campaniense-Maastrichtiense) de Valencia [PhD dissertation]. Valencia: Universidad de Valencia.Company, J., & Szentesi, Z. (2012). Amphibians from the Late Cretaceous Sierra Perenchiza Formation of the Chera Basin, Valencia Province, Spain. Cretaceous Research, 37, 240-245. doi:10.1016/j.cretres.2012.04.003Csiki-Sava, Z., Buffetaut, E., Ősi, A., Pereda-Suberbiola, X., & Brusatte, S. L. (2015). Island life in the Cretaceous - faunal composition, biogeography, evolution, and extinction of land-living vertebrates on the Late Cretaceous European archipelago. ZooKeys, 469, 1-161. doi:10.3897/zookeys.469.8439Erben, H. K., Hoefs, J., & Wedepohl, K. H. (1979). Paleobiological and isotopic studies of eggshells from a declining dinosaur species. Paleobiology, 5(4), 380-414. doi:10.1017/s0094837300016900García, R. A. (2007). An «egg-tooth»–like structure in titanosaurian sauropod embryos. Journal of Vertebrate Paleontology, 27(1), 247-252. doi:10.1671/0272-4634(2007)27[247:aesits]2.0.co;2Garcia, G., & Vianey-Liaud, M. (2001). Dinosaur eggshells as biochronological markers in Upper Cretaceous continental deposits. Palaeogeography, Palaeoclimatology, Palaeoecology, 169(1-2), 153-164. doi:10.1016/s0031-0182(01)00215-2Grellet-Tinner, G., Chiappe, L. M., & Coria, R. (2004). Eggs of titanosaurid sauropods from the Upper Cretaceous of Auca Mahuevo (Argentina). Canadian Journal of Earth Sciences, 41(8), 949-960. doi:10.1139/e04-049Grigorescu, D., Garcia, G., Csiki, Z., Codrea, V., & Bojar, A.-V. (2010). Uppermost Cretaceous megaloolithid eggs from the Haţeg Basin, Romania, associated with hadrosaur hatchlings: Search for explanation. Palaeogeography, Palaeoclimatology, Palaeoecology, 293(3-4), 360-374. doi:10.1016/j.palaeo.2010.03.031Izquierdo LA, Montero D, Pérez G, Urién V, Meijide M. 2001. Macroestructura de huevos de dinosaurios en el Cretácico superior de “La Rosaca” (Burgos, España). Actas de las I Jornadas Internacionales Sobre Paleontología de Dinosaurios y su Entorno. Ed. Colectivo Arqueológico y Paleontológico de Salas. Salas de los Infantes. p. 389–395.Jackson FD. 2007. Titanosaur reproductive biology: comparison of the Auca Mahuevo Titanosaur nesting locality (Argentina), to the Pinyes Megaloolithus nesting locality (Spain) [PhD dissertation]. Bozeman (MT): Montana State University.Jackson, F. D., Garrido, A., Schmitt, J. G., Chiappe, L. M., Dingus, L., & Loope, D. B. (2004). Abnormal, multilayered titanosaur (Dinosauria: Sauropoda) eggs from in situ clutches at the Auca Mahuevo locality, Neuquen Province, Argentina. Journal of Vertebrate Paleontology, 24(4), 913-922. doi:10.1671/0272-4634(2004)024[0913:amtdse]2.0.co;2Jackson, F. D., Varricchio, D. J., Jackson, R. A., Vila, B., & Chiappe, L. M. (2008). Comparison of water vapor conductance in a titanosaur egg from the Upper Cretaceous of Argentina and a Megaloolithus siruguei egg from Spain. Paleobiology, 34(2), 229-246. doi:10.1666/0094-8373(2008)034[0229:cowvci]2.0.co;2López-Martı́nez, N., Moratalla, J. J., & Sanz, J. L. (2000). Dinosaurs nesting on tidal flats. Palaeogeography, Palaeoclimatology, Palaeoecology, 160(1-2), 153-163. doi:10.1016/s0031-0182(00)00063-8Mohabey, D. M. (1998). Systematics of Indian Upper Cretaceous dinosaur and chelonian eggshells. Journal of Vertebrate Paleontology, 18(2), 348-362. doi:10.1080/02724634.1998.10011063Moratalla JJ. 1993. Restos indirectos de dinosaurios del registro español: paleoicnología de la Cuenca de (Jurásico superior-Cretácico inferior) y paleoología del Cretácico superior [PhD dissertation]. Madrid: Universidad Autónoma de Madrid.Moreno-Azanza, M., Bauluz, B., Canudo, J. I., Gasca, J. M., & Torcida Fernández-Baldor, F. (2016). Combined Use of Electron and Light Microscopy Techniques Reveals False Secondary Shell Units in Megaloolithidae Eggshells. PLOS ONE, 11(5), e0153026. doi:10.1371/journal.pone.0153026Moreno-Azanza, M., Bauluz, B., Canudo, J. I., Puértolas-Pascual, E., & Sellés, A. G. (2013). A re-evaluation of aff. Megaloolithidae eggshell fragments from the uppermost Cretaceous of the Pyrenees and implications for crocodylomorph eggshell structure. Historical Biology, 26(2), 195-205. doi:10.1080/08912963.2013.786067Oms, O., Dinarès-Turell, J., Vicens, E., Estrada, R., Vila, B., Galobart, À., & Bravo, A. M. (2007). Integrated stratigraphy from the Vallcebre Basin (southeastern Pyrenees, Spain): New insights on the continental Cretaceous−Tertiary transition in southwest Europe. Palaeogeography, Palaeoclimatology, Palaeoecology, 255(1-2), 35-47. doi:10.1016/j.palaeo.2007.02.039Ortega, F., Bardet, N., Barroso-Barcenilla, F., Callapez, P. M., Cambra-Moo, O., Daviero- Gómez, V., … Sanz, J. L. (2015). The biota of the Upper Cretaceous site of «Lo Hueco» (Cuenca, Spain). Journal of Iberian Geology, 41(1). doi:10.5209/rev_jige.2015.v41.n1.48657Rasskin-Gutman, D., Elez, J., Esteve-Altava, B., & López-Martínez, N. (2020). Reconstruction of the internal structure of the pore system of a complex dinosaur eggshell (Megaloolithus siruguei). Spanish Journal of Palaeontology, 28(1), 61. doi:10.7203/sjp.28.1.17831Riera, V., Oms, O., Gaete, R., & Galobart, À. (2009). The end-Cretaceous dinosaur succession in Europe: The Tremp Basin record (Spain). Palaeogeography, Palaeoclimatology, Palaeoecology, 283(3-4), 160-171. doi:10.1016/j.palaeo.2009.09.018Sellés, A. G., Bravo, A. M., Delclòs, X., Colombo, F., Martí, X., Ortega-Blanco, J., … Galobart, À. (2013). Dinosaur eggs in the Upper Cretaceous of the Coll de Nargó area, Lleida Province, south-central Pyrenees, Spain: Oodiversity, biostratigraphy and their implications. Cretaceous Research, 40, 10-20. doi:10.1016/j.cretres.2012.05.004Tanaka, K., & Zelenitsky, D. K. (2014). Comparisons between experimental and morphometric water vapor conductance in the eggs of extant birds and crocodiles: implications for predicting nest type in dinosaurs. Canadian Journal of Zoology, 92(12), 1049-1058. doi:10.1139/cjz-2014-0078Vianey-Liaud, M., Khosla, A., & Garcia, G. (2003). Relationships between European and Indian dinosaur eggs and eggshells of the oofamily Megaloolithidae. Journal of Vertebrate Paleontology, 23(3), 575-585. doi:10.1671/0272-4634(2003)023[0575:rbeaid]2.0.co;2Vianey-Liaud, M., & Lopez-Martinez, N. (1997). Late Cretaceous dinosaur eggshells from the Tremp Basin, southern Pyrenees, Lleida, Spain. Journal of Paleontology, 71(6), 1157-1171. doi:10.1017/s002233600003609xVila, B., Galobart, À., Canudo, J. I., Le Loeuff, J., Dinarès-Turell, J., Riera, V., … Gaete, R. (2012). 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Aristotelian Essentialism: Essence in the Age of Evolution

The advent of contemporary evolutionary theory ushered in the eventual decline of Aristotelian Essentialism (Æ) – for it is widely assumed that essence does not, and cannot have any proper place in the age of evolution. This paper argues that this assumption is a mistake: if Æ can be suitably evolved, it need not face extinction. In it, I claim that if that theory’s fundamental ontology consists of dispositional properties, and if its characteristic metaphysical machinery is interpreted within the framework of contemporary evolutionary developmental biology, an evolved essentialism is available. The reformulated theory of Æ offered in this paper not only fails to fall prey to the typical collection of criticisms, but is also independently both theoretically and empirically plausible. The paper contends that, properly understood, essence belongs in the age of evolution

Pere Alberch's developmental morphospaces and the evolution of cognition

In this article we argue for an extension of Pere Alberch's notion of developmental morphospace into the realm of cognition and introduce the notion of cognitive phenotype as a new tool for the evolutionary and developmental study of cognitive abilities