Analysis of variability in vertebral morphology and growth ring counts in two Carcharhinid sharks

Inter- and intra-regional variations in vertebrae morphology and growth increment counts (band counts) were analyzed for two carcharhinid shark species, Carcharhinus plumbeus (n = 10) and C. limbatus (n = 11). Five sequential vertebrae were removed from the cervical region, above the branchial chamber and posterior to the chondrocrainium, and thoracic region, below the first dorsal fin. Dorsal–ventral height, medial–lateral breadth, and caudal–cranial length were measured for each sampled vertebra. Results indicate no significant difference in vertebral morphology within a sampled region of the vertebral column. However, a significant difference in vertebral morphology was noted between regions for both shark species, with thoracic vertebrae consistently larger than cervical vertebrae. A sub-set of three vertebrae was taken from each sampled region of each shark for sectioning and counting of growth increments. Analyses of growth increment counts by two readers indicated no significant difference in band counts within and between sampled regions.     

Sex differences in the vertebral column of gombe chimpanzees

Male and female chimpanzees from Gombe National Park, Tanzania (Pan troglodytes schweinfurthii) differ in live body weights but not in cranial capacity or fore-and hindlimb long bone lengths. Skeletal dimensions of the limbs and vertebral column indicate a mosaic of sex differences. Vertebral column measurements generally are greater in males. While linear measurements identify differences in the breadth and depth of the lower cervical and upper thoracic vertebrae, areal assessments show significant differences in weight-bearing surfaces throughout the thoracic and lumbar segments. These can be interpreted in terms of distribution of weight and body composition (i.e. amount of musculature).     

Variation in number of vertebrae and gill rakers of sockeye salmon,Oncorhynchus nerka,in North America

Synopsis The latitudinal, regional, and annual variation in number of vertebrae and number of gill rakers present in sockeye salmon, Oncorhynchus nerka, stocks in North America was examined. Stocks in more northern areas had higher numbers of vertebrae and gill rakers than did those in more southern ones. Significant annual variability in the frequencies of these meristic characters within stocks was observed. When stocks were grouped into three regions (southern, central, and northern), heterogeneity in vertebral and gill raker frequencies was greater among regions than among stocks within the regions. Similarly, heterogeneity was greater among stocks than among sampling years within stocks. Differences in vertebral and gill raker frequencies are only useful for stock identification of sockeye salmon on a broad regional basis.     

Brief communication: Transportation and trauma: Dog‐sledding and vertebral compression in Alaskan Eskimos

Vertebral compression, as evidenced by compression of the centrum, was observed within two Native Alaskan skeletal samples. Information was collected from 1,071 and 656 vertebrae from Golovin Bay and Nunivak Island, Alaska, respectively. In addition, patterns of compression related vertebral change in each collection were characterized by sex and location within the vertebral column. The overall frequencies of vertebral compression were 3.6% (n = 721) at Golovin Bay and 1.7% (n = 403) at Nunivak Island for all observable thoracic and lumbar vertebrae (T1–L5). There was no statistically significant difference in the occurrence of compression among adults between these two collections. When examining the thoracic and lumbar vertebral segments by sex, females at Golovin Bay (4.5%; n = 442) exhibited a significantly higher frequency of vertebral compression than females at Nunivak (1.0%; n = 203). However, this difference in occurrence of compression could be accounted for by the age distributions of the two samples. No difference was noted between the males of the two collections. Compression frequencies in both samples are discussed in relation to the modes of transportation historically utilized by each community. Am J Phys Anthropol, 2010. © 2010 Wiley‐Liss, Inc.     

Regional variation in morphology of vertebral centra and intervertebral joints in striped bass,Morone saxatilis

The vertebral column of fishes has traditionally been divided into just two distinct regions, abdominal and caudal. Recently, however, developmental, morphological, and mechanical investigations have brought this traditional regionalization scheme into question. Alternative regionalization schema advocate the division of the abdominal vertebrae into cervical, abdominal, and in some cases, transitional regions. Here, we investigate regional variation at the level of the vertebrae and intervertebral joint (IVJ) tissues in the striped bass, Morone saxatilis. We use gross dissection, histology, and polarized light imaging to quantify vertebral height, width, length, IVJ length, IVJ tissue volume and cross‐sectional area, and vertical septum fiber populations, and angles of insertion. Our results reveal regional differences between the first four (most rostral) abdominal vertebrae and IVJs and the next six abdominal vertebrae and IVJs, supporting the recognition of a distinct cervical region. We found significant variation in vertebral length, width, and height from cranial to caudal. In addition, we see a significant decline in the volume of notochordal cells and the cross‐sectional area of the fibrous sheath from cranial to caudal. Further, polarized light imaging revealed four distinct fiber populations within the vertical septum in the cervical and abdominal regions in contrast with just one fiber population found in the caudal region. Measurement of the insertion angles of these fiber populations revealed significant differences between the cervical and abdominal regions. Differences in vertebral, IVJ, and vertical septum morphology all predict greater range of motion and decreased stiffness in the caudal region of the fish compared with the cervical and abdominal regions. J. Morphol., 2012. © 2011 Wiley Periodicals, Inc.     

Heritability and genetic correlation of abdominal and caudal vertebral numbers in latitudinal populations of the medaka <Emphasis Type="Italic">Oryzias latipes</Emphasis>

Body axes of fishes consist of two anatomically distinct types of vertebrae: abdominal and caudal. In the medaka Oryzias latipes, the number of abdominal vertebrae increases with increasing latitudes, whereas caudal vertebrae do not vary systematically across latitudes, suggesting local adaptation in abdominal vertebral numbers. However, because heritable variation in abdominal and caudal vertebral numbers has not been examined within each latitudinal population, it is not clear whether abdominal and caudal vertebrae can evolve independently. Offspring-midparent regression demonstrated substantial heritability of abdominal vertebral numbers in each of two latitudinal populations whereas the heritability of caudal vertebral numbers was not significant. Full-sib analyses revealed that intra-family variation was larger in caudal vertebrae than in abdominal vertebrae, indicating larger non-additive genetic variation and/or larger errors of development in the former. Moreover, the genetic correlation between abdominal and caudal vertebral numbers was very weak. These results suggest that abdominal and caudal vertebrae are controlled by separate developmental modules, which supports their independent evolution with local adaptation of abdominal vertebral numbers in this fish. On the other hand, the weak heritability of caudal vertebrae suggests that the evolution of caudal vertebrae may be restricted, causing unequal evolutionary lability between abdominal and caudal regions.     

Differential segmental growth of the vertebral column of the rat (Rattus norvegicus)

Despite the pervasive occurrence of segmental morphologies in the animal kingdom, the study of segmental growth is almost entirely lacking, but may have significant implications for understanding the development of these organisms. We investigate the segmental and regional growth of the entire vertebral column of the rat (Rattus norvegicus) by fitting a Gompertz curve to length and age data for each vertebra and each vertebral region. Regional lengths are calculated by summing constituent vertebral lengths and intervertebral space lengths for cervical, thoracic, lumbar, sacral, and caudal regions. Gompertz curves allow for the estimation of parameters representing neonatal and adult vertebral and regional lengths, as well as initial growth rate and the rate of exponential growth decay. Findings demonstrate differences between neonatal and adult rats in terms of relative vertebral lengths, and differential growth rates between sequential vertebrae and vertebral regions. Specifically, relative differences in the length of vertebrae indicate increasing differences caudad. Vertebral length in neonates increases from the atlas to the middle of the thoracic series and decreases in length caudad, while adult vertebral lengths tend to increase caudad. There is also a general trend of increasing vertebral and regional initial growth and rate of growth decay caudad. Anteroposterior patterns of growth are sexually dimorphic, with males having longer vertebrae than females at any given age. Differences are more pronounced (a) increasingly caudad along the body axis, and (b) in adulthood than in neonates. Elucidated patterns of growth are influenced by a combination of developmental, functional, and genetic factors.     

Functional anatomy and adaptation of the third to sixth thoracic vertebrae in primates using three-dimensional geometric morphometrics

The morphology of the cranial thoracic vertebrae has long been neglected in the study of primate skeletal functional morphology. This study explored the characteristics of the third to sixth thoracic vertebrae among various positional behavioural primates. A total of 67 skeletal samples from four species of hominoids, four of cercopithecoids, and two of platyrrhines were used. Computed tomography images of the thoracic vertebrae were converted to a three-dimensional (3D) bone surface, and 104 landmarks were obtained on the 3D surface. For size-independent shape analysis, the vertebrae were scaled to the same centroid size, and the normalised landmarks were registered using the generalised Procrustes method. Principle components of shape variation among samples were clarified using the variance–covariance matrix of the Procrustes residuals. The present study revealed that the transverse processes were more dorsally positioned in hominoids compared to non-hominoids. The results showed that not only a dorsolaterally oriented but also a dorsally positioned transverse process in relation to the vertebral arch contribute to the greater dorsal depth in hominoids than in monkeys. The thoracic vertebrae of Ateles and Nasalis show relatively dorsoventrally low and craniocaudally long vertebrae with craniocaudally long zygapophyses and craniocaudally long base/short tip of the caudally oriented spinous process, accompanied by a laterally oriented and craniocaudally long base of the transverse process. Despite being phylogenetically separated, the vertebral features of Ateles (suspensory platyrrhine with its prehensile tail's aid) are similar to those of Nasalis (arboreal quadrupedal/jumping/arm-swing colobine). The morphology of the third to sixth thoracic vertebrae tends to reflect the functional adaptation in relation to positional behaviour rather than the phylogenetic characteristics of hominoids, cercopithecoids, and platyrrhines.

     

Patterns of growth in the presacral vertebral column of the leopard gecko (Eublepharis macularius)

Postnatal growth patterns within the vertebral column may be informative about body proportions and regionalization. We measured femur length, lengths of all pre‐sacral vertebrae, and lengths of intervertebral spaces, from radiographs of a series of 21 Eublepharis macularius, raised under standard conditions and covering most of the ontogenetic body size range. Vertebrae were grouped into cervical, sternal, and dorsal compartments, and lengths of adjacent pairs of vertebrae were summed before analysis. Femur length was included as an index of body size. Principal component analysis of the variance‐covariance matrix of these data was used to investigate scaling among them. PC1 explained 94.19% of total variance, interpreted as the variance due to body size. PC1 differed significantly from the hypothetical isometric vector, indicating overall allometry. The atlas and axis vertebrae displayed strong negative allometry; the remainder of the vertebral pairs exhibited weak negative allometry, isometry or positive allometry. PC1 explained a markedly smaller amount of variance for the vertebral pairs of the cervical compartment than for the remainder of the vertebral pairs, with the exception of the final pair. The relative standard deviations of the eigenvalues from the PCAs of the three vertebral compartments indicated that the vertebrae of the cervical compartment were less strongly integrated by scaling than were the sternal or dorsal vertebrae, which did not differ greatly between themselves in their strong integration, suggesting that the growth of the cervical vertebrae is constrained by the mechanical requirements of the head. Regionalization of the remainder of the vertebral column is less clearly defined but may be associated with wave form propagation incident upon locomotion, and by locomotory changes occasioned by tail autotomy and regeneration. Femur length exhibits negative allometry relative to individual vertebral pairs and to vertebral column length, suggesting a change in locomotor requirements over the ontogenetic size range.     

A method for estimation of lateral and vertical mobility of platycoelous vertebrae of tetrapods

A new method for the estimation of dorsoventral and lateral mobility of platycoelous vertebrae with V-shaped (radial) articular facets on the zygapophyses is developed. This vertebral pattern is observed in dinosaurs, some other fossil reptiles, and in the cervical and lumbar regions of mammals. Based on theoretical biomechanical analysis of the intervertebral discs and articulations between zygapophyses, the estimation formulas are developed and calibrated, using precise measurements of mobility between cervical vertebrae of domestic sheep. The method is applied to the presacral vertebrae of the horned dinosaur Protoceratops andrewsi. In its cervical, lumbar, and anterior thoracic regions, the differences between the calculated amplitudes of movements and the sought true values are expected to range within ±5°. As compared to the sheep, Protoceratops shows a greater lateral mobility in the presacral region and reduced vertical mobility in the cervical region.     

Morphology of the vertebral centra in dolphins from the southwestern South Atlantic: A 3D morphometric approach and functional implications

In dolphins, centrum shape is one of the features that allows determination of stable and flexible regions in the vertebral column. The Commerson's (Cephalorhynchus commersonii; n = 37), Peale's (Lagenorhynchus australis; n = 24), dusky (Lagenorhynchus obscurus; n = 29), and hourglass dolphins (Lagenorhynchus cruciger; n = 10) are closely related species inhabiting the Southern Hemisphere that have diverse prey and habitat preferences. We applied 3D geometric morphometrics to describe differences in centrum shape along the vertebral columns of these species, and hypothesize how these differences may affect swimming. On each column, we chose a maximum of eight vertebrae and digitized 18 landmarks on each centrum with a Microscribe G2X. We explored shape differences amongst regions employing principal components analyses and computing Mahalanobis distances. We describe differences in centrum shape in relation to functional regions and among species; and analyze shape changes in relation to particular biomechanical requirements. The species studied here may be partially sympatric in the Southern Hemisphere, but they have important differences in foraging ecology and habitat preferences that could be related to differences in centrum shape along the vertebral column.     

Functional morphology of the primate head and neck

The vertebral column plays a key role in maintaining posture, locomotion, and transmitting loads between body components. Cervical vertebrae act as a bridge between the torso and head and play a crucial role in the maintenance of head position and the visual field. Despite its importance in positional behaviors, the functional morphology of the cervical region remains poorly understood, particularly in comparison to the thoracic and lumbar sections of the spinal column. This study tests whether morphological variation in the primate cervical vertebrae correlates with differences in postural behavior. Phylogenetic generalized least-squares analyses were performed on a taxonomically broad sample of 26 extant primate taxa to test the link between vertebral morphology and posture. Kinematic data on primate head and neck postures were used instead of behavioral categories in an effort to provide a more direct analysis of our functional hypothesis. Results provide evidence for a function-form link between cervical vertebral shape and postural behaviors. Specifically, taxa with more pronograde heads and necks and less kyphotic orbits exhibit cervical vertebrae with longer spinous processes, indicating increased mechanical advantage for deep nuchal musculature, and craniocaudally longer vertebral bodies and more coronally oriented zygapophyseal articular facets, suggesting an emphasis on curve formation and maintenance within the cervical lordosis, coupled with a greater resistance to translation and ventral displacement. These results not only document support for functional relationships in cervical vertebrae features across a wide range of primate taxa, but highlight the utility of quantitative behavioral data in functional investigations. Am J Phys Anthropol 156:531–542, 2015. © 2015 Wiley Periodicals, Inc.     

Variation of vertebral number in juvenile Oncorhynchus mykiss in relation to upstream distance from the ocean

Vertebral counts in fishes are set early in development by a complex interaction of genetic and environmental factors. Recently, R. M. McDowall showed that vertebral counts can differ between diadromous and non-diadromous galaxiids, and suggested similar life history-based variation might also occur in the closely related Salmonidae. We examined variation in vertebral counts of juvenile rainbow trout, Oncorhynchus mykiss, from within a single large basin where anadromous and non-anadromous forms co-occur. Juveniles were collected at 29 sites within the Klamath River basin using a nested sampling design. Mean vertebral counts varied among regions, among complexes within regions, and among sites within complexes. Regression showed that mean vertebral counts decreased with increasing distance from the ocean. This decrease could not be explained by differences in temperature within the basin during early development. However, the decrease is consistent with the expectation that non-anadromous forms have fewer vertebrae and are more common at sites distant from the ocean, and suggests that further study of the relationship between vertebral counts and life history form in O. mykiss is warranted.

Phenotypic integration of the cervical vertebrae in the Hominoidea (Primates)

Phenotypic integration and modularity represent important factors influencing evolutionary change. The mammalian cervical vertebral column is particularly interesting in regards to integration and modularity because it is highly constrained to seven elements, despite widely variable morphology. Previous research has found a common pattern of integration among quadrupedal mammals, but integration patterns also evolve in response to locomotor selective pressures like those associated with hominin bipedalism. Here, I test patterns of covariation in the cervical vertebrae of three hominoid primates (Hylobates, Pan, Homo) who engage in upright postures and locomotion. Patterns of integration in the hominoid cervical vertebrae correspond generally to those previously found in other mammals, suggesting that integration in this region is highly conserved, even among taxa that engage in novel positional behaviors. These integration patterns reflect underlying developmental as well as functional modules. The strong integration between vertebrae suggests that the functional morphology of the cervical vertebral column should be considered as a whole, rather than in individual vertebrae. Taxa that display highly derived morphologies in the cervical vertebrae are likely exploiting these integration patterns, rather than reorganizing them. Future work on vertebrates without cervical vertebral number constraints will further clarify the evolution of integration in this region.     

Segmental relationship between somites and vertebral column in zebrafish

The segmental heritage of all vertebrates is evident in the character of the vertebral column. And yet, the extent to which direct translation of pattern from the somitic mesoderm and de novo cell and tissue interactions pattern the vertebral column remains a fundamental, unresolved issue. The elements of vertebral column pattern under debate include both segmental pattern and anteroposterior regional specificity. Understanding how vertebral segmentation and anteroposterior positional identity are patterned requires understanding vertebral column cellular and developmental biology. In this study, we characterized alignment of somites and vertebrae, distribution of individual sclerotome progeny along the anteroposterior axis and development of the axial skeleton in zebrafish. Our clonal analysis of zebrafish sclerotome shows that anterior and posterior somite domains are not lineage-restricted compartments with respect to distribution along the anteroposterior axis but support a 'leaky' resegmentation in development from somite to vertebral column. Alignment of somites with vertebrae suggests that the first two somites do not contribute to the vertebral column. Characterization of vertebral column development allowed examination of the relationship between vertebral formula and expression patterns of zebrafish Hox genes. Our results support co-localization of the anterior expression boundaries of zebrafish hoxc6 homologs with a cervical/thoracic transition and also suggest Hox-independent patterning of regionally specific posterior vertebrae.     

Morphology and Function of the Vertebral Column in Remingtonocetus domandaensis (Mammalia, Cetacea) from the Middle Eocene Domanda Formation of Pakistan

The archaeocete family Remingtonocetidae is a group of early cetaceans known from the Eocene of India and Pakistan. Previous studies of remingtonocetids focused primarily on cranial anatomy due to a paucity of well-preserved postcranial material. Here we describe the morphology of the known vertebral column in Remingtonocetus domandaensis based largely on a single well-preserved partial skeleton recovered from the upper Domanda Formation of Pakistan. The specimen preserves most of the precaudal vertebral column in articulation and includes seven complete cervical vertebrae, ten partial to complete thoracic vertebrae, six complete lumbar vertebrae, and the first three sacral vertebrae. Cervical centra are long and possess robust, imbricating transverse processes that stabilized the head and neck. Lumbar vertebrae allowed for limited flexibility and probably served primarily to stabilize the lumbar column during forceful retraction of the hind limbs. Vertebral evidence, taken together with pelvic and femoral morphology, is most consistent with interpretation of Remingtonocetus domandaensis as an animal that swam primarily by powerful movement of its hind limbs rather than dorsoventral undulation of its body axis.     

Evolution of the axial skeleton in armadillos (Mammalia,Dasypodidae)

Intraspecific and interspecific variation in cervical, thoracic, and lumbar region of the vertebral column of Dasypodidae were examined in a phylogenetic framework. The number of vertebrae for each region were recorded for 86 specimens and metric data for each vertebra (centrum length, high, and width) were recorded for 72 specimens, including eight of the nine living genera. The number of vertebrae and degree of fusion between them were used to define four characters which were plotted on two alternative phylogenies of Dasypodidae. The ratio between centrum height and width is similar across all taxa analyzed except for Chlamyphorus, which exhibits a deviation in the last two lumbars. Tolypeutes matacus is unique among the taxa examined in having a second co-osified bone called postcervical bone, which is a fusion of the seventh cervical and first thoracic vertebrae. The thoraco-lumbar numbers of dasypodids are reduced when compared with other xenarthrans and are more diverse than those of some other mammalian clades of similar geological age and higher ecomorphological diversity. Changes in size are somewhat coupled with changes in the number of body segments. Independent of the phylogenetic framework taken, changes in size are accompanied with small changes in numbers of thoracolumbar vertebrae within each genus. There are functional and phylogenetic correlates for changes in number of thoraco-lumbar vertebrae in dasypodids.     

GEOGRAPHIC VARIATION AND PHENOTYPIC PLASTICITY OF NUMBER OF TRUNK VERTEBRAE IN SLENDER SALAMANDERS,BATRACHOSEPS (CAUDATA: PLETHODONTIDAE)

To understand the evolutionary significance of geographic variation, one must identify the factors that generate phenotypic differences among populations. I examined the causes of geographic variation in and evolutionary history of number of trunk vertebrae in slender salamanders, Batrachoseps (Caudata: Plethodontidae). Number of trunk vertebrae varies at many taxonomic levels within Batrachoseps. Parallel clines in number occur along an environmental gradient in three lineages in the Coast Ranges of California. These parallel clines may signal either adaptation or a shared phenotypically plastic response to the environmental gradient. By raising eggs from 10 populations representing four species of Batrachoseps, I demonstrated that number of trunk vertebrae can be altered by the developmental temperature; however, the degree of plasticity is insufficient to account for geographic variation. Thus, the geographic variation results largely from genetic variation. Number of trunk vertebrae covaries with body size and shape in diverse vertebrate taxa, including Batrachoseps. I hypothesize that selection for different degrees of elongation, possibly related to fossoriality, has led to the extensive evolution of number of trunk vertebrae in Batrachoseps. Analysis of intrapopulational variation revealed sexual dimorphism in both body shape and number of trunk vertebrae, but no correlation between these variables in either sex. Females are more elongate than males, a pattern that has been attributed to fecundity selection in other taxa. Patterns of covariation among different classes of vertebrae suggest that some intrapopulational variation in number results from changes in vertebral identity rather than changes in segmentation.     

Morphological variation of the thoracolumbar vertebrae in Macropodidae and its functional relevance

The present study was designed to investigate how the form of the marsupial thoracolumbar vertebrae varies to cope with the particular demands of diverse loading and locomotor behaviors. The vertebral columns of 10 species of Macropodidae, with various body masses and modes of locomotion, together with two other arboreal marsupials, koala and cuscus, were selected. Seventy-four three-dimensional landmark coordinates were acquired on each of the 10 last presacral vertebrae of the 70 vertebral columns. The interspecific variations of the third lumbar vertebra (L3, which approximates the mean) and the transitional patterns of the thoracolumbar segments were examined using the combined approach of generalized Procrustes analysis (GPA) and principal components analysis (PCA). The results of analyses of an individual vertebra (L3) and of the transitional patterns indicate significant interspecific differences. In the L3 study the first PC shows allometric shape variation, while the second PC seems to relate to adaptation for terrestrial versus arboreal locomotion. When the L3 vertebrae of the common spotted cuscus and koala are included for comparison, the vertebra of the tree kangaroo occupies an intermediate position between the hopping kangaroo and these arboreal marsupials. The L3 vertebrae in the arboreal marsupials possess a distinct dorsoventrally expanded vertebral body, and perpendicularly orientated spinous and transverse processes. The results of the present study suggest that vertebral shape in the kangaroo and wallaroos provides a structural adaptation to hopping through a relatively enlarged loading area and powerful lever system. In contrast, the small-sized bettongs (or rat kangaroos) have a relatively flexible column and elongated levers for the action of back muscles that extend and laterally flex the spine. The complex pattern of vertebral shape transition in the last 10 presacral vertebrae was examined using PCAs that compare between species information about vertebral shape variation along the thoracolumbar column. The results reinforce and emphasize important aspects of the patterns of variation seen in the detailed analysis of the third lumbar vertebra. The results also imply that size, spinal loading pattern, and locomotor behavior exert an influence on shaping the vertebra. Further, the morphological adaptations are consistent within these marsupials and this opens up the possibility that this kind of analysis may be useful in making functional inferences from fossil material.     

Building an evolutionary innovation: Differential growth in the modified vertebral elements of the zebrafish Weberian apparatus

The Weberian apparatus, a complex assemblage of greatly modified vertebral elements, significantly enhances hearing within Otophysi. Ultimately we are interested in investigating the genetic mechanisms responsible for the origin, development and morphological diversification of these vertebral elements in the Weberian apparatus of otophysan fishes. However, a necessary first step involves identifying changes in growth of this region as compared with the vertebrae from which these modified elements purportedly derive. Using an ontogenetic series of the zebrafish, Danio rerio, we collected growth data for specific elements within the Weberian apparatus, including neural arches, ribs, and parapophyses. These data are compared to both serially homologous structures in posterior thoracic vertebrae (which act as internal controls) and vertebral elements from the same axial levels in three other non-otophysan teleosts. Significant differences in growth rate were found among serially homologous structures, as well as at equivalent axial levels in different species. Uniform changes in growth rates (in which all structures derived from a specific somite were equally affected) were not found, suggesting precise targeting of morphological change to specific structures. The variation in growth of anterior vertebrae in and among species was greater than expected. This variation in growth rates created developmental patterns unique to each species. Such patterns of growth may help illuminate the specific heterochronic mechanisms required for the origin and subsequent morphological diversification of the Weberian apparatus. This morphological diversity is exemplified by the multitude of forms seen in the cypriniform Weberian apparatus. Understanding patterns of growth in discrete elements of the Weberian apparatus allows us to hypothesize as to the specific developmental changes, likely constituting differences in gene expression in pathways involved in bone and cartilage differentiation, responsible for this morphological diversity.