Studies in Quantitative Paleontology. III. An Application of Sequential Analysis to the Comparison of Growth Stages and Growth Series

The albertosaurines Albertosaurus sarcophagus and Gorgosaurus libratus are among the best represented tyrannosaurids, known from nearly complete growth series. These specimens provide an opportunity to study mandibular biomechanical properties and tooth morphology in order to infer changes in feeding behavior and bite force through ontogeny in tyrannosaurids. Mandibular force profiles reveal that the symphyseal region of albertosaurines is consistently stronger in bending than the middentary region, indicating that the anterior extremity of the jaws played an important role in prey capture and handling through ontogeny. The symphyseal region was better adapted to withstand torsional stresses than in most non-avian theropods, but not to the extent seen in Tyrannosaurus rex, suggesting that albertosaurine feeding behavior may have involved less bone crushing or perhaps relatively smaller prey than in T. rex. The constancy of these biomechanical properties at all known growth stages indicates that although albertosaurines maintained a similar feeding strategy through ontogeny, prey size/type had to change between juvenile and mature individuals. This ontogenetic dietary shift likely happened when individuals reached a mandibular length of ~58 cm, a size at which teeth shift from ziphodont to incrassate in shape and bite force begins to increase exponentially. The fact that large albertosaurines were capable of generating bite forces equivalent to similar-sized tyrannosaurines suggests that no significant differences in jaw closing musculature existed between the two clades and that the powerful bite of T. rex is the result of its large body size rather than of unique adaptations related to a specialized ecology.

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Ontogeny and relationships of the Ordovician odontopleurid trilobite Ceratocara, with new species from Argentina and New York

Journal of Paleontology ◽

10.1017/s0022336000039007 ◽

1997 ◽

Vol 71 (1) ◽

pp. 108-125 ◽

Cited By ~ 21

Author(s):

Brian D. E. Chatterton ◽

Gregory D. Edgecombe ◽

Norberto E. Vaccari ◽

Beatriz G. Waisfeld

Keyword(s):

New York ◽

Phylogenetic Analysis ◽

New Species ◽

San Juan ◽

Growth Stages ◽

Upper Ordovician ◽

Growth Series ◽

Middle Ordovician ◽

Argentine Precordillera ◽

A New Species

A growth series is described for a new species of Ceratocara Ramsköld, 1991, C. argentina, from the Upper Ordovician (Caradoc) part of the Las Aguaditas Formation, near Jáchal in San Juan Province, Argentine Precordillera. Another new species of Ceratocara, C. shawi, is described from the Middle Ordovician (lower Chazy) Crown Point Formation of New York. A phylogenetic analysis is presented for these species, other well-known Ordovician species of Ceratocara, and some Ordovician species of Ceratocephala, with Ceratocephalina tridens Whittington, 1956, as outgroup. The analysis presented supports the monophyly of both Ceratocara and Ceratocephala Warder, 1838, their divergence having occurred by the middle Arenig. The ontogenies of the Ceratocara species from Argentina and New York, complete from protaspid to holaspid growth stages, are some of the best preserved odontopleurid ontogenies described to date. Material of Ceratocephala triacantheis Whittington and Evitt, 1954, from the Crown Point Formation of the Chazy Group of New York, is discriminated from that of Ceratocara shawi.

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Larvae and relationships of the Calymenina (Trilobita)

Journal of Paleontology ◽

10.1017/s0022336000018424 ◽

1990 ◽

Vol 64 (2) ◽

pp. 255-277 ◽

Cited By ~ 48

Author(s):

B. D. E. Chatterton ◽

D. J. Siveter ◽

G. D. Edgecombe ◽

A. S. Hunt

Keyword(s):

Cladistic Analysis ◽

Data Matrix ◽

Growth Stages ◽

Low Latitude ◽

Growth Series ◽

Larval Stages ◽

Close Relationship ◽

Life History Patterns ◽

Devonian Age ◽

Relationship Of

Up to four discrete protaspid larval stages are described for calymenid trilobites of Ordovician to Devonian age. The earliest growth stages are nonadult-like planktonic protaspides; later protaspides are adult-like and benthonic. In contrast, the related homalonotid trilobites apparently lack planktonic protaspides, but have up to two large benthonic protaspid stages that are similar in form to calymenid benthonic protaspides. These differences in life history patterns between these families are reflected in their paleobiogeographic distributions. Calymenids werre widely dispersed from Ordovician to Devonian times, both being common in warm, low latitude provinces (particularly from the Late Ordovician onwards) and well represented in cooler, higher latitude regions. The paleogeographic distribution of the homalonotids during the Ordovician (Arenig to the Ashgill) was concentrated in high paleolatitudes, with only a few forms occurring at low paleolatitudes (often in deeper, cooler environments?). Both families survived the Ordovician–Silurian mass extinction, with the calymenids again being widely dispersed but the homalonotids being best represented in the cool-water Malvinokaffric Province and in other regions where they are largely restricted to clastic facies.So few complete growth series of calymenine trilobites are known that it is unlikely that the ontogenies of taxa that form parts of ancestor–descendant clades can be identified. However, some evidence for heterochronic, particularly paedomorphic (neotenic), evolution is suggested for larval stages of members of both the Calymenidae and the Homalonotidae. Such possible neotenic evolution leading to very large planktonic larval stages of calymenid trilobites during the Devonian could have enhanced dispersal during a period of widespread warm and equable climates. Comparisons of homalonotid protaspides with equivalent stages of calymenids support the close relationship of these families within the Calymenina. A data matrix based upon characters of protaspides of two calymenine trilobites (Flexicalymene Shirley, 1936, and Brongniartella Reed, 1918) and eight other trilobites, belonging to the Phacopina (Calyptaulax), Cheirurina (Physemataspis and Hyrokybe), Proetida (Scharyia), Lichida (Acanthopyge), Odontopleurida (Diacanthaspis), Corynexochida (Bathyuriscus), and Ptychopariida (Crassifimbra) was subjected to cladistic analysis using the parsimony program “Hennig 86.” The shortest length cladogram produced is consistent with the inclusion of the Homalonotidae in the Calymenina, and inclusion of the Calymenina in the order Phacopida. “Cheirurina” is the paraphyletic “stem group” of Phacopina. The hypothesis that Lonchocephalidae is most closely related to part of post-Cambrian Phacopida is poorly supported by protaspid characters.

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Ontogeny of the crinoid Eucalyptocrinites

Journal of Paleontology ◽

10.1017/s0022336000061679 ◽

1968 ◽

Vol 42 (S2) ◽

pp. 99-118 ◽

Cited By ~ 6

Author(s):

Donald B. Macurda

Keyword(s):

Regression Analysis ◽

External Surface ◽

Growth Parameters ◽

Correlation Coefficients ◽

Growth Stages ◽

Specific Difference ◽

Growth Series ◽

Similar Mode ◽

Evolutionary Success ◽

Mode Of Development

The calyx of a Paleozoic camerate crinoid is composed of a mosaic of large, polygonal plates. Ontogenetic size increase results from accretionary additions of calcite to thecal plates. When new calcite is added to the lateral edge of a plate, a new layer is also added over the external surface, obscuring the earlier growth stages of the plate. However, ontogenetic development can be studied by measurement of a growth series. This is illustrated by regression analysis of growth series of two Silurian species of the genus Eucalyptocrinites: E. crassus (Hall) and E. tuberculatus (Miller & Dyer). Growth of the principal plates is isometric; most growth parameters have a very similar mode of development in both species; only a few parameters show any specific difference. A table of correlation coefficients for 51 growth parameters demonstrates that the entire development of the calyx was highly coordinated throughout life, with coefficients almost always equal to or larger than 0.90. Integrated expansion of the cup plates provided a larger cavity for the viscera. The stem increased in diameter to support the increasingly larger theca, as did the diameter of the root system. Growth of the vaulted plates in the upper part of the theca provided a protective recess for the arms, which probably contributed to the evolutionary success of this animal on a widespread basis during the Silurian and Devonian.Based upon growth studies, the following synonymies are suggested: E. constrictus Hall, E. ellipticus Miller, E. ovalis Hall, and E. subglobosus Miller are synonyms of E. crassus; E. clrodi Miller is a synonym of E. tuberculatus Miller & Dyer.

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Major cranial changes during Triceratops ontogeny

Proceedings of The Royal Society B Biological Sciences ◽

10.1098/rspb.2006.3643 ◽

2006 ◽

Vol 273 (1602) ◽

pp. 2757-2761 ◽

Cited By ~ 70

Author(s):

John R Horner ◽

Mark B Goodwin

Keyword(s):

Older Adults ◽

Posterior Margin ◽

Sexual Maturity ◽

Shape Function ◽

Early Ontogeny ◽

Foramen Magnum ◽

Growth Stages ◽

Growth Series ◽

Visual Identity ◽

Suture Fusion

This is the first cranial ontogenetic assessment of Triceratops , the well-known Late Cretaceous dinosaur distinguished by three horns and a massive parietal–squamosal frill. Our analysis is based on a growth series of 10 skulls, ranging from a 38 cm long baby skull to about 2 m long adult skulls. Four growth stages correspond to a suite of ontogenetic characters expressed in the postorbital horns, frill, nasal, epinasal horn and epoccipitals. Postorbital horns are straight stubs in early ontogeny, curve posteriorly in juveniles, straighten in subadults and recurve anteriorly in adults. The posterior margin of the baby frill is deeply scalloped. In early juveniles, the frill margin becomes ornamented by 17–19 delta-shaped epoccipitals. Epoccipitals are dorsoventrally compressed in subadults, strongly compressed and elongated in adults and ultimately merge onto the posterior frill margin in older adults. Ontogenetic trends within and between growth stages include: posterior frill margin transitions from scalloped to wavy and smooth; progressive exclusion of the supraoccipital from the foramen magnum; internal hollowing at the base of the postorbital horns; closure of the midline nasal suture; fusion of the epinasal onto the nasals; and epinasal expansion into a morphologically variable nasal horn. We hypothesize that the changes in horn orientation and epoccipital shape function to allow visual identity of juveniles, and signal their attainment of sexual maturity.

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A high-resolution growth series of Tyrannosaurus rex obtained from multiple lines of evidence

PeerJ ◽

10.7717/peerj.9192 ◽

2020 ◽

Vol 8 ◽

pp. e9192 ◽

Cited By ~ 1

Author(s):

Thomas D. Carr

Keyword(s):

New Technology ◽

Cladistic Analysis ◽

High Growth ◽

Morphological Characters ◽

High Growth Rate ◽

Chronological Age ◽

Growth Stages ◽

Ontogenetic Changes ◽

Data Set ◽

Growth Series

Background During the growth of complex multicellular organisms, chronological age, size and morphology change together in a hierarchical and coordinated pattern. Among extinct species, the growth of Tyrannosaurus rex has received repeated attention through quantitative analyses of relative maturity and chronological age. Its growth series shows an extreme transformation from shallow skulls in juveniles to deep skulls in adults along with a reduction in tooth count, and its growth curve shows that T. rex had a high growth rate in contrast to its closest relatives. However, separately, these sets of data provide an incomplete picture of the congruence between age, size, and relative maturity in this exemplar species. The goal of this work is to analyze these data sets together using cladistic analysis to produce a single hypothesis of growth that includes all of the relevant data. Methods The three axes of growth were analyzed together using cladistic analysis, based on a data set of 1,850 morphological characters and 44 specimens. The analysis was run in TNT v.1.5 under a New Technology search followed by a Traditional search. Correlation tests were run in IBM SPSS Statistics v. 24.0.0.0. Results An initial analysis that included all of the specimens recovered 50 multiple most parsimonious ontograms a series of analyses identified 13 wildcard specimens. An analysis run without the wildcard specimens recovered a single most parsimonious tree (i.e., ontogram) of 3,053 steps. The ontogram is composed of 21 growth stages, and all but the first and third are supported by unambiguously optimized synontomorphies. T. rex ontogeny can be divided into five discrete growth categories that are diagnosed by chronological age, morphology, and, in part, size (uninformative among adults). The topology shows that the transition from shallow to deep skull shape occurred between 13 and 15 years of age, and the size of the immediate relatives of T. rex was exceeded between its 15th and 18th years. Although size and maturity are congruent among juveniles and subadults, congruence is not seen among adults; for example, one of the least mature adults (RSM 2523.8) is also the largest and most massive example of the species. The extreme number of changes at the transition between juveniles and subadults shows that the ontogeny of T. rex exhibits secondary metamorphosis, analogous to the abrupt ontogenetic changes that are seen at sexual maturity among teleosts. These results provide a point of comparison for testing the congruence between maturity and chronological age, size, and mass, as well as integrating previous work on functional morphology into a rigorous ontogenetic framework. Comparison of the growth series of T. rex with those of outgroup taxa clarifies the ontogenetic trends that were inherited from the common ancestor of Archosauriformes.

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A histological survey of avian post-natal skeletal ontogeny

PeerJ ◽

10.7717/peerj.12160 ◽

2021 ◽

Vol 9 ◽

pp. e12160

Author(s):

Jessie Atterholt ◽

Holly N. Woodward

Keyword(s):

Body Size ◽

Bone Growth ◽

High Growth ◽

Growth Stages ◽

Crown Group ◽

Bony Tissue ◽

Growth Series ◽

Developmental Mode ◽

Evolutionary Origins ◽

Osteological Development

Bone histology of crown-group birds is a research topic of great interest, permitting insight into the evolution of remarkably high growth rates in this clade and variation across the altricial-precocial spectrum. In this study, we describe microanatomical characteristics of the humerus and femur in partial growth series from 14 crown group birds representing ten major clades (Struthioniformes, Galliformes, Apodiformes, Columbiformes, Charadriiformes, Accipitriformes, Strigiformes, Psittaciformes, Falconiformes, and Passeriformes). Our goals were to: (1) describe the microanatomy of each individual; (2) make inter-and intra-taxonomic comparisons; (3) assess patterns that correspond with developmental mode; and (4) to further parse out phylogenetic, developmental, and functional constraints on avian osteological development. Across taxa, the femoral and humeral tissue of neonates can be broadly characterized as highly-vascularized, disorganized woven bone with great variation in cortical thickness (inter-and intrataxonomically, within an individual specimen, and within a single section). The tissue of precocial chicks is relatively more mature at hatching than in altricial, but other categories along the developmental spectrum were less easy to distinguish, thus we were unable to identify a definitive histological proxy for developmental mode. We did not find evidence to support hypotheses that precocial chicks exclusively have thicker cortices and more mature bone in the femur than the humerus at time of hatching; instead, this is a characteristic of nearly all taxa (regardless of developmental mode), suggesting deep evolutionary origins and the effects of developmental channeling. Bone tissue in adults exhibited unexpected variation, corresponding to differences in body size. Large-bodied birds have cortices of fibrolamellar bone, but organization of tissue increases and vascularity decreases with diminishing body size. The outer circumferential layer (OCL) also appears at earlier growth stages in small-bodied taxa. Thus, while the OCL is indicative of a cessation of appositional growth it is not always indicative of cortical maturity (that is, maximum organization of bony tissue for a given taxon). Small size is achieved by truncating the period of fast growth; manipulation of the timing of offset of bone growth is therefore an important factor in changing growth trajectories to alter adult body size.

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Neuron-glia relationship during the early postembryonic development of the nervous system in some oligochaetes

Proceedings, annual meeting, Electron Microscopy Society of America ◽

10.1017/s0424820100083874 ◽

1978 ◽

Vol 36 (2) ◽

pp. 600-601

Author(s):

Wiktor Djaczenko ◽

Carmen Calenda Cimmino

Keyword(s):

Nervous System ◽

Glial Cells ◽

Early Period ◽

Postembryonic Development ◽

Ruthenium Red ◽

The Body ◽

Tubifex Tubifex ◽

Growth Stages ◽

Cell Processes ◽

Cytoplasmic Processes

The simplicity of the developing nervous system of oligochaetes makes of it an excellent model for the study of the relationships between glia and neurons. In the present communication we describe the relationships between glia and neurons in the early periods of post-embryonic development in some species of oligochaetes.Tubifex tubifex (Mull. ) and Octolasium complanatum (Dugès) specimens starting from 0. 3 mm of body length were collected from laboratory cultures divided into three groups each group fixed separately by one of the following methods: (a) 4% glutaraldehyde and 1% acrolein fixation followed by osmium tetroxide, (b) TAPO technique, (c) ruthenium red method.Our observations concern the early period of the postembryonic development of the nervous system in oligochaetes. During this period neurons occupy fixed positions in the body the only observable change being the increase in volume of their perikaryons. Perikaryons of glial cells were located at some distance from neurons. Long cytoplasmic processes of glial cells tended to approach the neurons. The superimposed contours of glial cell processes designed from electron micrographs, taken at the same magnification, typical for five successive growth stages of the nervous system of Octolasium complanatum are shown in Fig. 1. Neuron is designed symbolically to facilitate the understanding of the kinetics of the growth process.

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Characterisation of multilayer materials using a position-sensitive atom probe

Proceedings, annual meeting, Electron Microscopy Society of America ◽

10.1017/s0424820100176253 ◽

1990 ◽

Vol 48 (4) ◽

pp. 624-625

Author(s):

RAD Mackenzie ◽

G D W Smith ◽

A. Cerezo ◽

J A Liddle ◽

CRM Grovenor ◽

...

Keyword(s):

Quantum Well ◽

Quantum Wells ◽

Spatial Information ◽

Chemical Vapour ◽

Atom Probe ◽

Organic Chemical ◽

Growth Stages ◽

Multiple Quantum ◽

Quartz Wool ◽

Position Sensitive

The position sensitive atom probe (POSAP), described briefly elsewhere in these proceedings, permits both chemical and spatial information in three dimensions to be recorded from a small volume of material. This technique is particularly applicable to situations where there are fine scale variations in composition present in the material under investigation. We report the application of the POSAP to the characterisation of semiconductor multiple quantum wells and metallic multilayers.The application of devices prepared from quantum well materials depends on the ability to accurately control both the quantum well composition and the quality of the interfaces between the well and barrier layers. A series of metal organic chemical vapour deposition (MOCVD) grown GaInAs-InP quantum wells were examined after being prepared under three different growth conditions. These samples were observed using the POSAP in order to study both the composition of the wells and the interface morphology. The first set of wells examined were prepared in a conventional reactor to which a quartz wool baffle had been added to promote gas intermixing. The effect of this was to hold a volume of gas within the chamber between growth stages, leading to a structure where the wells had a composition of GalnAsP lattice matched to the InP barriers, and where the interfaces were very indistinct. A POSAP image showing a well in this sample is shown in figure 1. The second set of wells were grown in the same reactor but with the quartz wool baffle removed. This set of wells were much better defined, as can be seen in figure 2, and the wells were much closer to the intended composition, but still with measurable levels of phosphorus. The final set of wells examined were prepared in a reactor where the design had the effect of minimizing the recirculating volume of gas. In this case there was again further improvement in the well quality. It also appears that the left hand side of the well in figure 2 is more abrupt than the right hand side, indicating that the switchover at this interface from barrier to well growth is more abrupt than the switchover at the other interface.

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Cytokinins from terminal buds of Euphoria longana during different growth stages

Physiologia Plantarum ◽

10.1034/j.1399-3054.1997.990125.x ◽

1997 ◽

Vol 99 (1) ◽

pp. 185-189

Author(s):

Wen-Shaw Chen ◽

Kuang-Liang Huang ◽

Hsiao-Ching Yu

Keyword(s):

Growth Stages ◽

Terminal Buds ◽

Different Growth Stages

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