Haversian canals

@article{kornblum1964the,
    author = "KORNBLUM, S. SANFORD and KELLY, PATRICK J.",
    title = "The Lacunae and Haversian Canals in Tibial Cortical Bone from Ischemic and Non-Ischemic Limbs",
    year = "1964",
    journal = "The Journal of Bone \& Joint Surgery",
    url = "https://doi.org/10.2106/00004623-196446040-00009",
    doi = "10.2106/00004623-196446040-00009",
    number = "4",
    openalex = "W2239344049",
    pages = "797-810",
    volume = "46"
}

@article{bouvier1977dinosaur,
    author = "Bouvier, Marianne",
    title = "DINOSAUR HAVERSIAN BONE AND ENDOTHERMY",
    year = "1977",
    journal = "Evolution",
    url = "https://doi.org/10.1111/j.1558-5646.1977.tb01028.x",
    doi = "10.1111/j.1558-5646.1977.tb01028.x",
    number = "2",
    openalex = "W2325766925",
    pages = "449-450",
    volume = "31",
    references = "bakker1972anatomical, doi1010160021929074900645, doi101017s0021859600040491, doi101242jcss310361111, doi1021060000462319705208000005, openalexw2247544283"
}

@article{muto1980crystalloid,
    author = "MUTO, HIROSHI and YOSHIOKA, IKUO",
    title = "Crystalloid Substance and Closed Haversian Canals in the Compact Bone Tissue of Human Bone",
    year = "1980",
    journal = "Okajimas Folia Anatomica Japonica",
    url = "https://doi.org/10.2535/ofaj1936.57.2-3\_115",
    doi = "10.2535/ofaj1936.57.2-3\_115",
    number = "2-3",
    openalex = "W2003535940",
    pages = "115-127",
    volume = "57",
    references = "doi1010160003996962901255, doi101111j174966321955tb40054x, doi1021060000462319573901000015, doi1023073570764, openalexw1529872137, openalexw2623776990"
}

@article{nelson1980counts1,
    author = "Nelson, R. C. and Bookhout, T. A",
    title = "Counts of periosteal layers invalid for aging Canada Geese",
    year = "1980",
    journal = "Journal of Wildlife Management, v. 44, p. 518-521",
    note = "talkorigins\_source = {true}; raw\_reference = {Nelson, R. C., and Bookhout, T. A., 1980, Counts of periosteal layers invalid for aging Canada Geese: Journal of Wildlife Management, v. 44, p. 518-521.}"
}

@incollection{ricqles1980tissue3,
    author = "Ricqles, A. R",
    editor = "Thomas, D. K. and Olson, E. C.",
    title = "Tissue Structures of Dinosaur Bones: Functional Significance and Possible Relation to Dinosaur Physiology",
    year = "1980",
    booktitle = "A Cold Look at the Warm Blooded Dinosaurs",
    publisher = "Washington, D.C., American Association for the Advancement of Science, p. 103-140",
    note = "talkorigins\_source = {true}; raw\_reference = {Ricqles, A. R., 1980, Tissue Structures of Dinosaur Bones: Functional Significance and Possible Relation to Dinosaur Physiology, in Thomas, D. K., and Olson, E. C., eds., A Cold Look at the Warm Blooded Dinosaurs: Washington, D.C., American Association for the Advancement of Science, p. 103-140.}"
}

Patterns of bone loss in the axial and the appendicular skeleton were studied in 185 normal volunteers (105 women and 82 men; age range, 20--89 yr) and in 76 women and 9 men with vertebral fractures due to osteoporosis. Bone mineral density was measured in vivo at the lumbar spine (predominantly trabecular bone) by dual photon absorptiometry and at the midradius (greater than 95% cortical bone) and distal radius (75% cortical and 25% trabecular bone) by single photon absorptiometry. In normal women, bone diminution from the vertebrae began in young adulthood and was linear. In the appendicular skeleton, bone diminution did not occur until age 50 yr, was accelerated from aged 51 to 65 yr, and then decelerated somewhat after age 65 yr. Overall bone diminution throughout life was 47% for the vertebrae, 30% for the midradius, and 39% for the distal radius. In normal men, vertebral and appendicular bone diminution with aging was minimal or insignificant. Mean bone mineral density was lower in patients with osteoporosis than in age- and sex-matched normal subjects at all three scanning sites, although spinal measurements discriminated best; however, there was considerable overlap. By age 65 yr, half of the normal women (and by age 85 yr, virtually all of them) had vertebral bone mineral density values below the 90th percentile of women with vertebral fractures and, thus, might be considered to have asymptomatic osteoporosis. For men, the degree of overlap was less. The data suggest that disproportionate loss of trabecular bone from the axial skeleton is a distinguishing characteristic of spinal osteoporosis.

@article{doi101172jci110039,
    author = "Riggs, B. Lawrence and Wahner, Heinz W. and Dunn, W L and Mazess, Richard B. and Offord, Kenneth P. and Melton, L. Joseph",
    title = "Differential changes in bone mineral density of the appendicular and axial skeleton with aging: relationship to spinal osteoporosis.",
    year = "1981",
    journal = "Journal of Clinical Investigation",
    abstract = "Patterns of bone loss in the axial and the appendicular skeleton were studied in 185 normal volunteers (105 women and 82 men; age range, 20--89 yr) and in 76 women and 9 men with vertebral fractures due to osteoporosis. Bone mineral density was measured in vivo at the lumbar spine (predominantly trabecular bone) by dual photon absorptiometry and at the midradius (greater than 95\% cortical bone) and distal radius (75\% cortical and 25\% trabecular bone) by single photon absorptiometry. In normal women, bone diminution from the vertebrae began in young adulthood and was linear. In the appendicular skeleton, bone diminution did not occur until age 50 yr, was accelerated from aged 51 to 65 yr, and then decelerated somewhat after age 65 yr. Overall bone diminution throughout life was 47\% for the vertebrae, 30\% for the midradius, and 39\% for the distal radius. In normal men, vertebral and appendicular bone diminution with aging was minimal or insignificant. Mean bone mineral density was lower in patients with osteoporosis than in age- and sex-matched normal subjects at all three scanning sites, although spinal measurements discriminated best; however, there was considerable overlap. By age 65 yr, half of the normal women (and by age 85 yr, virtually all of them) had vertebral bone mineral density values below the 90th percentile of women with vertebral fractures and, thus, might be considered to have asymptomatic osteoporosis. For men, the degree of overlap was less. The data suggest that disproportionate loss of trabecular bone from the axial skeleton is a distinguishing characteristic of spinal osteoporosis.",
    url = "https://doi.org/10.1172/jci110039",
    doi = "10.1172/jci110039",
    openalex = "W2122745505"
}

We measured bone mineral density (BMD) of the proximal femur, lumbar spine, or both by dual photon absorptiometry in 205 normal volunteers (123 women and 82 men; age range 20 to 92 yr) and in 31 patients with hip fractures (26 women and 5 men; mean age, 78 yr). For normal women, the regression of BMD on age was negative and linear at each site; overall decrease during life was 58% in the femoral neck, 53% in the intertrochanteric region of the femur, and 42% in the lumbar spine. For normal men, the age regression was linear also; the rate of decrease in BMD was two-thirds of that in women for femoral neck and intertrochanteric femur but was only one-fourth of that in women for lumbar spine. This difference may explain why the female/male ratio is 2:1 for hip fractures but 8:1 for vertebral fractures. The standard deviation (Z-score) from the sex-specific age-adjusted normal mean in 26 women with hip fracture averaged -0.31 (P < 0.05) for the femoral neck, -0.53 (P < 0.01) for the intertrochanteric femur, and +0.24 (NS) for the lumbar spine; results were similar for 5 men with hip fractures. By contrast, for 27 additional women, ages 51-65 yr, with only nontraumatic vertebral fractures, the Z-score was -1.92 (P < 0.001) for the lumbar spine. Thus, contrary to the view that osteoporosis is a single age-related entity, our data suggest the existence of two distinct syndromes. One form, "postmenopausal osteoporosis," is characterized by excessive and disproportionate trabecular bone loss, involves a small subset of women in the early postmenopausal period, and is associated mainly with vertebral fractures. The other form, "senile osteoporosis," is characterized by proportionate loss of both cortical and trabecular bone, involves essentially the entire population of aging women and, to a lesser extent, aging men, and is associated with hip fractures or vertebral fractures or both.

@article{doi101172jci110667,
    author = "Riggs, B. Lawrence and Wahner, Heinz W. and Seeman, Ego and Offord, Kenneth P. and Dunn, W. L. and Mazess, R. B. and Johnson, Kenneth A. and Melton, L. Joseph",
    title = "Changes in Bone Mineral Density of the Proximal Femur and Spine with Aging",
    year = "1982",
    journal = "Journal of Clinical Investigation",
    abstract = {We measured bone mineral density (BMD) of the proximal femur, lumbar spine, or both by dual photon absorptiometry in 205 normal volunteers (123 women and 82 men; age range 20 to 92 yr) and in 31 patients with hip fractures (26 women and 5 men; mean age, 78 yr). For normal women, the regression of BMD on age was negative and linear at each site; overall decrease during life was 58\% in the femoral neck, 53\% in the intertrochanteric region of the femur, and 42\% in the lumbar spine. For normal men, the age regression was linear also; the rate of decrease in BMD was two-thirds of that in women for femoral neck and intertrochanteric femur but was only one-fourth of that in women for lumbar spine. This difference may explain why the female/male ratio is 2:1 for hip fractures but 8:1 for vertebral fractures. The standard deviation (Z-score) from the sex-specific age-adjusted normal mean in 26 women with hip fracture averaged -0.31 (P < 0.05) for the femoral neck, -0.53 (P < 0.01) for the intertrochanteric femur, and +0.24 (NS) for the lumbar spine; results were similar for 5 men with hip fractures. By contrast, for 27 additional women, ages 51-65 yr, with only nontraumatic vertebral fractures, the Z-score was -1.92 (P < 0.001) for the lumbar spine. Thus, contrary to the view that osteoporosis is a single age-related entity, our data suggest the existence of two distinct syndromes. One form, "postmenopausal osteoporosis," is characterized by excessive and disproportionate trabecular bone loss, involves a small subset of women in the early postmenopausal period, and is associated mainly with vertebral fractures. The other form, "senile osteoporosis," is characterized by proportionate loss of both cortical and trabecular bone, involves essentially the entire population of aging women and, to a lesser extent, aging men, and is associated with hip fractures or vertebral fractures or both.},
    url = "https://doi.org/10.1172/jci110667",
    doi = "10.1172/jci110667",
    openalex = "W1826010384"
}

@inproceedings{reid1984the2,
    author = "Reid, R. E. H",
    title = "The histology of dinosaurian bone, and its possible bearing on dinosaur physiology",
    year = "1984",
    booktitle = "Symposium of the Zoological Society, London, v. 52, p. 629-663",
    note = "talkorigins\_source = {true}; raw\_reference = {Reid, R. E. H., 1984, The histology of dinosaurian bone, and its possible bearing on dinosaur physiology: Symposium of the Zoological Society, London, v. 52, p. 629-663.}"
}

We studied the morphology of the haversian canals in the osteopenic cortical bone of the medial femoral neck from patients with rheumatoid arthritis and compared the findings with those in patients with osteoarthritis and with uncomplicated coxa valga. In the rheumatoid bone, the diameters of the canals were larger and many more contained osteoclasts. Fewer haversian canals showed only lining cells than in the osteoarthritic or coxa valga patients. In bone from rheumatoid patients, especially in canals with osteoclasts, small blood vessels were frequently lined by tall endothelial cells with an infiltration of mononuclear cells. These morphological differences are discussed with reference to the possible mechanisms of loss of cortical bone in rheumatoid arthritis and other conditions.

@article{takashima1989inflammatory,
    author = "Takashima, T and Kawai, K and Hirohata, K and Miki, A and Mizoguti, H and Cooke, TD",
    title = "Inflammatory cell changes in haversian canals. A possible cause of osteoporosis in rheumatoid arthritis",
    year = "1989",
    journal = "The Journal of Bone and Joint Surgery. British volume",
    abstract = "We studied the morphology of the haversian canals in the osteopenic cortical bone of the medial femoral neck from patients with rheumatoid arthritis and compared the findings with those in patients with osteoarthritis and with uncomplicated coxa valga. In the rheumatoid bone, the diameters of the canals were larger and many more contained osteoclasts. Fewer haversian canals showed only lining cells than in the osteoarthritic or coxa valga patients. In bone from rheumatoid patients, especially in canals with osteoclasts, small blood vessels were frequently lined by tall endothelial cells with an infiltration of mononuclear cells. These morphological differences are discussed with reference to the possible mechanisms of loss of cortical bone in rheumatoid arthritis and other conditions.",
    url = "https://doi.org/10.1302/0301-620x.71b4.2768320",
    doi = "10.1302/0301-620x.71b4.2768320",
    number = "4",
    openalex = "W2336808046",
    pages = "671-676",
    volume = "71-B"
}

ABSTRACT The bone microstructure of three third metatarsals and two tibiae of differing sizes from Troodon formosus was examined. Bone in T. formosus passed through three ontogenetic stages: rapid fibro-lamellar, moderate lamellar-zonal, and slow avascular lamellar growth. Highly-vascularized fibro-lamellar bone accounted for the majority of growth, with adult size possibly being reached in 3 to 5 years. No significant growth occurred after T. formosus reached a body weight of roughly 50 kg (femur: tibia length 310:425 mm). Abundant vasculature together with dense Haversian bone suggests T. formosus had a relatively high metabolism. Other gross morphological features (large brain, large eyes, cursorial adaptations including high tibia: femur ratio) would indicate T. formosus was very active and possibly endothermic.

@article{doi10108002724634199310011490,
    author = "Varricchio, David J.",
    title = "Bone microstructure of the Upper Cretaceous theropod dinosaur Troodon formosus",
    year = "1993",
    journal = "Journal of Vertebrate Paleontology",
    abstract = "ABSTRACT The bone microstructure of three third metatarsals and two tibiae of differing sizes from Troodon formosus was examined. Bone in T. formosus passed through three ontogenetic stages: rapid fibro-lamellar, moderate lamellar-zonal, and slow avascular lamellar growth. Highly-vascularized fibro-lamellar bone accounted for the majority of growth, with adult size possibly being reached in 3 to 5 years. No significant growth occurred after T. formosus reached a body weight of roughly 50 kg (femur: tibia length 310:425 mm). Abundant vasculature together with dense Haversian bone suggests T. formosus had a relatively high metabolism. Other gross morphological features (large brain, large eyes, cursorial adaptations including high tibia: femur ratio) would indicate T. formosus was very active and possibly endothermic.",
    url = "https://doi.org/10.1080/02724634.1993.10011490",
    doi = "10.1080/02724634.1993.10011490",
    openalex = "W1993549596",
    references = "bouvier1977dinosaur, crossref1976allosaurus, doi1010160031018271900447, doi101038332256a0, doi101086410790, doi101111j146979981985tb04915x, doi1023071443592, doi1023073514834, openalexw575222456, russell1969a, spotila1973a, wilson1985stenonychosaurus"
}

Histological and ultrastructural evaluation of the ends of long bones of juvenile dinosaurs from the Upper Cretaceous Two Medicine Formation of Montana revealed the preservation of growth plates. Growth plates are discs of cartilage present near the ends of growing long bones that generate bone elongation. Comparison of the fossils with modern taxa demonstrated homology of the growth plate in birds and dinosaurs. The presence of an avian-type growth plate in dinosaurs adds a shared derived anatomical character corroborating inclusion of birds within the Dinosauria. Additionally, possession of a growth plate, which in birds is capable of producing rapid determinate long bone growth, implies that an avian developmental pattern may have been present in these dinosaurs.

@article{doi101126science26251422020,
    author = "Barreto, Claudia and Albrecht, Ralph M. and Bjorling, Dale E. and Horner, John R. and Wilsman, Norman J.",
    title = "Evidence of the Growth Plate and the Growth of Long Bones in Juvenile Dinosaurs",
    year = "1993",
    journal = "Science",
    abstract = "Histological and ultrastructural evaluation of the ends of long bones of juvenile dinosaurs from the Upper Cretaceous Two Medicine Formation of Montana revealed the preservation of growth plates. Growth plates are discs of cartilage present near the ends of growing long bones that generate bone elongation. Comparison of the fossils with modern taxa demonstrated homology of the growth plate in birds and dinosaurs. The presence of an avian-type growth plate in dinosaurs adds a shared derived anatomical character corroborating inclusion of birds within the Dinosauria. Additionally, possession of a growth plate, which in birds is capable of producing rapid determinate long bone growth, implies that an avian developmental pattern may have been present in these dinosaurs.",
    url = "https://doi.org/10.1126/science.262.5142.2020",
    doi = "10.1126/science.262.5142.2020",
    openalex = "W2007794110",
    references = "bouvier1977dinosaur, doi101002jor1100090306, doi101017s247526300000091x, doi101038248168a0, doi101038282296a0, doi101038297675a0, doi101038361064a0, doi1010970124139819921100000022, doi1021060000462319876902000002, doi1023071443592, openalexw575222456"
}

Twelve different bones from the skeleton of the holotype specimen of the hadrosaurian dinosaur Hypacrosaurus stebingeri were thin-sectioned to evaluate the significance of lines of arrested growth (LAGs) in age assessments. The presence of an external fundamental system (EFS) at the external surface of the cortex and mature epiphyses indicate that the Hypacrosaurus specimen had reached adulthood and growth had slowed considerably from earlier stages. The number of LAGs varied from none in the pedal phalanx to as many as eight in the tibia and femur. Most elements had experienced considerable Haversian reconstruction that had most likely obliterated many LAGs. The tibia was found to have experienced the least amount of reconstruction, but was still not optimal for skeletochronology because the LAGs were difficult to count near the periosteal surface. Additionally, the numbers of LAGs within the EFS vary considerably around the circumference of a single element and among elements. Counting LAGs from a single bone to assess skeletochronology appears to be unreliable, particularly when a fundamental system exists. Because LAGs are plesiomorphic for tetrapods, and because they are present in over a dozen orders of mammals, they have no particular physiological meaning that can be generalized to particular amniote groups without independent physiological evidence. Descriptions of dinosaur physiology as “intermediate” between the physiology of living reptiles and that of living birds and mammals may or may not be valid, but cannot be based reliably on the presence of LAGs.

@article{doi101017s0094837300021308,
    author = "Horner, John R. and de Ricqlès, Armand and Padian, Kevin",
    title = "Variation in dinosaur skeletochronology indicators: implications for age assessment and physiology",
    year = "1999",
    journal = "Paleobiology",
    abstract = "Twelve different bones from the skeleton of the holotype specimen of the hadrosaurian dinosaur Hypacrosaurus stebingeri were thin-sectioned to evaluate the significance of lines of arrested growth (LAGs) in age assessments. The presence of an external fundamental system (EFS) at the external surface of the cortex and mature epiphyses indicate that the Hypacrosaurus specimen had reached adulthood and growth had slowed considerably from earlier stages. The number of LAGs varied from none in the pedal phalanx to as many as eight in the tibia and femur. Most elements had experienced considerable Haversian reconstruction that had most likely obliterated many LAGs. The tibia was found to have experienced the least amount of reconstruction, but was still not optimal for skeletochronology because the LAGs were difficult to count near the periosteal surface. Additionally, the numbers of LAGs within the EFS vary considerably around the circumference of a single element and among elements. Counting LAGs from a single bone to assess skeletochronology appears to be unreliable, particularly when a fundamental system exists. Because LAGs are plesiomorphic for tetrapods, and because they are present in over a dozen orders of mammals, they have no particular physiological meaning that can be generalized to particular amniote groups without independent physiological evidence. Descriptions of dinosaur physiology as “intermediate” between the physiology of living reptiles and that of living birds and mammals may or may not be valid, but cannot be based reliably on the presence of LAGs.",
    url = "https://doi.org/10.1017/s0094837300021308",
    doi = "10.1017/s0094837300021308",
    openalex = "W2285964556",
    references = "chinsamy1994dinosaur, chinsamy1998polar, crossref1998encyclopedia, doi101002jmor1051080103, doi101017s0094837300013543, doi10108002724634199310011490, doi10108002724634199510011271, doi101093clinids222240, doi101111j109636422000tb00016x, doi101111j155856461974tb00777x, doi1016710272463420000200115lbhoth20co2, doi1023071564284, doi105860choice353642, openalexw424753225, openalexw575222456, openalexw597127060, openalexw648632191, openalexw991367939"
}

ABSTRACT Ontogenetic changes in the bone histology of Maiasaura peeblesorum are revealed by six relatively distinct but gradational growth stages: early and late nestling, early and late juvenile, sub-adult, and adult. These stages are distinguished not only by relative size but by changes in the histological patterns of bones at each stage. In general, the earliest stages are marked by spongy bone matrix with large vascular canals. Through growth, the cortical bone differentiates into fibro-lamellar tissue that tends to become more regularly layered in the outer cortex. By the sub-adult stage, lines of arrested growth (LAGs) begin to appear regularly. Resorption lines and substantial Haversian substitution in many long bones also begin to appear at this stage, and the external cortex has a lamellar-zonal structure in some bones that indicates imminent cessation of growth. Judging by the rates of apposition of similar bone tissues in living amniotes, and by the number and placement of LAGs, these patterns suggest that young Maiasaura nestlings grew at very high rates, and at high and moderately high rates during later nestling, juvenile, and sub-adult stages, slowing to low and very low growth rates in adults (7–9 m total length). The nesting period would have lasted one to two months, late juvenile size (3.5 meters) would have been reached in one or two years, and adult size in six to eight years, depending on the basis for extrapolating bone growth rates. The histological tissues, patterns, and inferred growth rates of the bones of Maiasaura are completely different from those of living non-avian reptiles, generally similar to those of most other dinosaurs and pterosaurs for which data are available, and much like those of extant birds and mammals. No living reptiles (except birds) grow to adult size at these rates, nor do they show these histological patterns. We conclude that Maiasaura did not grow at all like living non-avian reptiles, which cannot be considered informative models for most aspects of dinosaurian growth (or physiology, to the extent that growth rates reflect metabolism). The use of lines of arrested growth (LAGs) to infer dinosaurian physiology has never been tested and is not supported by independent lines of evidence; their use in calculating age is also more complex than previously suggested and should not be based on single bones.

@article{doi1016710272463420000200115lbhoth20co2,
    author = "Horner, John R. and de Ricqlès, Armand and Padian, Kevin",
    title = "Long bone histology of the hadrosaurid dinosaur Maiasaura peeblesorum: growth dynamics and physiology based on an ontogenetic series of skeletal elements",
    year = "2000",
    journal = "Journal of Vertebrate Paleontology",
    abstract = "ABSTRACT Ontogenetic changes in the bone histology of Maiasaura peeblesorum are revealed by six relatively distinct but gradational growth stages: early and late nestling, early and late juvenile, sub-adult, and adult. These stages are distinguished not only by relative size but by changes in the histological patterns of bones at each stage. In general, the earliest stages are marked by spongy bone matrix with large vascular canals. Through growth, the cortical bone differentiates into fibro-lamellar tissue that tends to become more regularly layered in the outer cortex. By the sub-adult stage, lines of arrested growth (LAGs) begin to appear regularly. Resorption lines and substantial Haversian substitution in many long bones also begin to appear at this stage, and the external cortex has a lamellar-zonal structure in some bones that indicates imminent cessation of growth. Judging by the rates of apposition of similar bone tissues in living amniotes, and by the number and placement of LAGs, these patterns suggest that young Maiasaura nestlings grew at very high rates, and at high and moderately high rates during later nestling, juvenile, and sub-adult stages, slowing to low and very low growth rates in adults (7–9 m total length). The nesting period would have lasted one to two months, late juvenile size (3.5 meters) would have been reached in one or two years, and adult size in six to eight years, depending on the basis for extrapolating bone growth rates. The histological tissues, patterns, and inferred growth rates of the bones of Maiasaura are completely different from those of living non-avian reptiles, generally similar to those of most other dinosaurs and pterosaurs for which data are available, and much like those of extant birds and mammals. No living reptiles (except birds) grow to adult size at these rates, nor do they show these histological patterns. We conclude that Maiasaura did not grow at all like living non-avian reptiles, which cannot be considered informative models for most aspects of dinosaurian growth (or physiology, to the extent that growth rates reflect metabolism). The use of lines of arrested growth (LAGs) to infer dinosaurian physiology has never been tested and is not supported by independent lines of evidence; their use in calculating age is also more complex than previously suggested and should not be based on single bones.",
    url = "https://doi.org/10.1671/0272-4634(2000)020[0115:lbhoth]2.0.co;2",
    doi = "10.1671/0272-4634(2000)020[0115:lbhoth]2.0.co;2",
    openalex = "W2179073245",
    references = "chinsamy1994dinosaur, chinsamy1998polar, doi101001jama195602970180082039, doi101002jmor1051080103, doi1010079781489957405, doi101017s0094837300012331, doi101017s0094837300013543, doi101017s0094837300021308, doi101029sc005p0175, doi101038282296a0, doi10108002724634199310011490, doi101093clinids222240, doi101126science26251422020, doi1016660094837320010270039coosea20co2, doi105962bhltitle113905, openalexw2259112626, openalexw648632191, openalexw991367939, reid1984primary"
}

For millions of years, clues to the biology of dinosaurs lie locked within the microscopic structure of their bones. Here one of the world's leading experts on fossil bone microstructure synthesizes more than 150 years of research to expose the meaning of dinosaur bone microstructure. In this first book dedicated to dinosaur bone microstructure, Anusuya Chinsamy-Turan begins with a general overview of living bone structure and composition and then explains how bone structure changes upon death and during fossilization. She meticulously unravels why the microscopic structure of fossil bone remains intact after millions of years of fossilization. She also provides a photographic atlas of the kinds of bone tissue found in dinosaurs and expounds on their biological significance. The final chapters offer insight into growth patterns of dinosaurs and the biology of Mesozoic birds. The book concludes with a reflective discussion on what bone microstructure can and cannot disclose about dinosaur physiology. Drawing from sources across the field of bone histology, Chinsamy-Turan paints a holistic view of the current state of the science and presents a fresh perspective on the relevance of the field to understanding the Dinosauria.

@article{doi105860choice432219,
    author = "Chinsamy-Turan, Anusuya 1962-",
    title = "The microstructure of dinosaur bone: deciphering biology with fine-scale techniques",
    year = "2005",
    journal = "Choice Reviews Online",
    abstract = "For millions of years, clues to the biology of dinosaurs lie locked within the microscopic structure of their bones. Here one of the world's leading experts on fossil bone microstructure synthesizes more than 150 years of research to expose the meaning of dinosaur bone microstructure. In this first book dedicated to dinosaur bone microstructure, Anusuya Chinsamy-Turan begins with a general overview of living bone structure and composition and then explains how bone structure changes upon death and during fossilization. She meticulously unravels why the microscopic structure of fossil bone remains intact after millions of years of fossilization. She also provides a photographic atlas of the kinds of bone tissue found in dinosaurs and expounds on their biological significance. The final chapters offer insight into growth patterns of dinosaurs and the biology of Mesozoic birds. The book concludes with a reflective discussion on what bone microstructure can and cannot disclose about dinosaur physiology. Drawing from sources across the field of bone histology, Chinsamy-Turan paints a holistic view of the current state of the science and presents a fresh perspective on the relevance of the field to understanding the Dinosauria.",
    url = "https://doi.org/10.5860/choice.43-2219",
    doi = "10.5860/choice.43-2219",
    openalex = "W651090911"
}

This review brings together a complex and extensive literature to address the question of whether it is possible to distinguish human from nonhuman bone using the histological appearance of cortical bone. The mammalian species included are rat, hare, badger, racoon dog, cat, dog, pig, cow, goat, sheep, deer, horse, water buffalo, bear, nonhuman primates, and human and are therefore not exhaustive, but cover those mammals that may contribute to a North American or Eurasian forensic assemblage. The review has demonstrated that differentiation of human from certain nonhuman species is possible, including small mammals exhibiting Haversian bone tissue and large mammals exhibiting plexiform bone tissue. Pig, cow, goat, sheep, horse, and water buffalo exhibit both plexiform and Haversian bone tissue and where only Haversian bone tissue exists in bone fragments, differentiation of these species from humans is not possible. Other primate Haversian bone tissue is also not distinguishable from humans. Where differentiation using Haversian bone tissue is undertaken, both the general microstructural appearance and measurements of histological structures should be applied. Haversian system diameter and Haversian canal diameter are the most optimal and diagnostic measurements to use. Haversian system density may be usefully applied to provide an upper and lower limit for humans.

@article{doi101111j15564029200600368x,
    author = "Hillier, Maria L. and Bell, Lynne",
    title = "Differentiating Human Bone from Animal Bone: A Review of Histological Methods",
    year = "2007",
    journal = "Journal of Forensic Sciences",
    abstract = "This review brings together a complex and extensive literature to address the question of whether it is possible to distinguish human from nonhuman bone using the histological appearance of cortical bone. The mammalian species included are rat, hare, badger, racoon dog, cat, dog, pig, cow, goat, sheep, deer, horse, water buffalo, bear, nonhuman primates, and human and are therefore not exhaustive, but cover those mammals that may contribute to a North American or Eurasian forensic assemblage. The review has demonstrated that differentiation of human from certain nonhuman species is possible, including small mammals exhibiting Haversian bone tissue and large mammals exhibiting plexiform bone tissue. Pig, cow, goat, sheep, horse, and water buffalo exhibit both plexiform and Haversian bone tissue and where only Haversian bone tissue exists in bone fragments, differentiation of these species from humans is not possible. Other primate Haversian bone tissue is also not distinguishable from humans. Where differentiation using Haversian bone tissue is undertaken, both the general microstructural appearance and measurements of histological structures should be applied. Haversian system diameter and Haversian canal diameter are the most optimal and diagnostic measurements to use. Haversian system density may be usefully applied to provide an upper and lower limit for humans.",
    url = "https://doi.org/10.1111/j.1556-4029.2006.00368.x",
    doi = "10.1111/j.1556-4029.2006.00368.x",
    openalex = "W1983236345",
    references = "doi101002ajpa1330230215, doi101016s0379073800003765"
}

Abstract Long bones (femora, humeri) are the most abundant remains of sauropod dinosaurs. Their length is a good proxy for body length and body mass, and their histology is informative about ontogenetic age. Here we provide a comparative assessment of histologic changes in growth series of several sauropod taxa, including diplodocids (Apatosaurus, Diplodocus, indeterminate Diplodocinae from the Tendaguru Beds and from the Morrison Formation), basal macronarians (Camarasaurus, Brachiosaurus, Europasaurus), and titanosaurs (Phuwiangosaurus, Ampelosaurus). A total of 167 long bones, mainly humeri and femora, and 18 limb girdle bones were sampled. Sampling was performed by core drilling at prescribed locations at midshaft, and 13 histologic ontogenetic stages (HOS stages) were recognized. Because growth of all sauropod long bones is quite uniform, with laminar fibrolamellar bone being the dominant tissue, HOS stages could be recognized across taxa, although with minor differences. Histologic ontogenetic stages generally correlate closely with body size and thus provide a means to resolve important issue like the ontogenetic status of questionable specimens. We hypothesize that sexual maturity was attained at HOS-8, well before maximum size was attained, but we did not find sexually differentiated growth trajectories subsequent to HOS-8. On the basis of HOS stages, we detected two morphotypes in the Camarasaurus sample, a small one (type 1) and a larger one (type 2), presumably representing different species or sexual dimorphism.

@article{doi1016660094837320080340247ositlb20co2,
    author = "Klein, Nicole and Sander, Martin",
    title = "Ontogenetic stages in the long bone histology of sauropod dinosaurs",
    year = "2008",
    journal = "Paleobiology",
    abstract = "Abstract Long bones (femora, humeri) are the most abundant remains of sauropod dinosaurs. Their length is a good proxy for body length and body mass, and their histology is informative about ontogenetic age. Here we provide a comparative assessment of histologic changes in growth series of several sauropod taxa, including diplodocids (Apatosaurus, Diplodocus, indeterminate Diplodocinae from the Tendaguru Beds and from the Morrison Formation), basal macronarians (Camarasaurus, Brachiosaurus, Europasaurus), and titanosaurs (Phuwiangosaurus, Ampelosaurus). A total of 167 long bones, mainly humeri and femora, and 18 limb girdle bones were sampled. Sampling was performed by core drilling at prescribed locations at midshaft, and 13 histologic ontogenetic stages (HOS stages) were recognized. Because growth of all sauropod long bones is quite uniform, with laminar fibrolamellar bone being the dominant tissue, HOS stages could be recognized across taxa, although with minor differences. Histologic ontogenetic stages generally correlate closely with body size and thus provide a means to resolve important issue like the ontogenetic status of questionable specimens. We hypothesize that sexual maturity was attained at HOS-8, well before maximum size was attained, but we did not find sexually differentiated growth trajectories subsequent to HOS-8. On the basis of HOS stages, we detected two morphotypes in the Camarasaurus sample, a small one (type 1) and a larger one (type 2), presumably representing different species or sexual dimorphism.",
    url = "https://doi.org/10.1666/0094-8373(2008)034[0247:ositlb]2.0.co;2",
    doi = "10.1666/0094-8373(2008)034[0247:ositlb]2.0.co;2",
    openalex = "W2177631336",
    references = "chinsamy1994dinosaur, doi101016jtree200508012, doi101029sc005p0175, doi101038nature04633, doi101046j10963642200200029x, doi101073pnas0708903105, doi10108002724634199310011490, doi101098rsbl20070254, doi101098rspb20042829, doi101111j109636422000tb02201x, doi1015159781400849505, doi1016660094837320000260466lhotts20co2, doi1016660094837320010270039coosea20co2, doi1016710272463420000200115lbhoth20co2, martinsander2006bone"
}

BACKGROUND: Sauropterygia is an abundant and successful group of Triassic marine reptiles. Phylogenetic relationships of Triassic Sauropterygia have always been unstable and recently questioned. Although specimens occur in high numbers, the main problems are rareness of diagnostic material from the Germanic Basin and uniformity of postcranial morphology of eosauropterygians. In the current paper, morphotypes of humeri along with their corresponding bone histologies for Lower to Middle Muschelkalk sauropterygians are described and interpreted for the first time in a phylogenetic context. METHODOLOGY/PRINCIPAL FINDINGS: Nothosaurus shows a typical plesiomorphic lamellar-zonal bone type, but varying growth patterns and the occurrence of a new humerus morphotype point to a higher taxonomic diversity than was known. In contrast to the enormous morphological variability of eosauropterygian humeri not assigned to Nothosaurus, their long bone histology is relatively uniform and can be divided into two histotypes. Unexpectedly, both of these histotypes reveal abundant fibrolamellar bone throughout the cortex. This pushes the origin of fibrolamellar bone in Sauropterygia back from the Cretaceous to the early Middle Triassic (early Anisian). Histotype A is assigned to Cymatosaurus, a basal member of the Pistosauroidea, which includes the plesiosaurs as derived members. Histotype B is related to the pachypleurosaur Anarosaurus. Contrary to these new finds, the stratigraphically younger pachypleurosaur Neusticosaurus shows the plesiomorphic lamellar-zonal bone type and an incomplete endochondral ossification, like Nothosaurus. CONCLUSIONS/SIGNIFICANCE: Histological results hypothetically discussed in a phylogenetical context have a large impact on the current phylogenetic hypothesis of Sauropterygia, leaving the pachypleurosaurs polyphyletic. On the basis of histological data, Neusticosaurus would be related to Nothosaurus, whereas Anarosaurus would follow the pistosaur clade. Furthermore, the presence of fibrolamellar bone, which is accompanied with increased growth rates and presumably even with increased metabolic rates, already in Anarosaurus and Cymatosaurus can explain the success of the Pistosauroidea, the only sauropterygian group to survive into the Jurassic and give rise to the pelagic plesiosaur radiation.

@article{doi101371journalpone0011613,
    author = "Klein, Nicole",
    title = "Long Bone Histology of Sauropterygia from the Lower Muschelkalk of the Germanic Basin Provides Unexpected Implications for Phylogeny",
    year = "2010",
    journal = "PLoS ONE",
    abstract = "BACKGROUND: Sauropterygia is an abundant and successful group of Triassic marine reptiles. Phylogenetic relationships of Triassic Sauropterygia have always been unstable and recently questioned. Although specimens occur in high numbers, the main problems are rareness of diagnostic material from the Germanic Basin and uniformity of postcranial morphology of eosauropterygians. In the current paper, morphotypes of humeri along with their corresponding bone histologies for Lower to Middle Muschelkalk sauropterygians are described and interpreted for the first time in a phylogenetic context. METHODOLOGY/PRINCIPAL FINDINGS: Nothosaurus shows a typical plesiomorphic lamellar-zonal bone type, but varying growth patterns and the occurrence of a new humerus morphotype point to a higher taxonomic diversity than was known. In contrast to the enormous morphological variability of eosauropterygian humeri not assigned to Nothosaurus, their long bone histology is relatively uniform and can be divided into two histotypes. Unexpectedly, both of these histotypes reveal abundant fibrolamellar bone throughout the cortex. This pushes the origin of fibrolamellar bone in Sauropterygia back from the Cretaceous to the early Middle Triassic (early Anisian). Histotype A is assigned to Cymatosaurus, a basal member of the Pistosauroidea, which includes the plesiosaurs as derived members. Histotype B is related to the pachypleurosaur Anarosaurus. Contrary to these new finds, the stratigraphically younger pachypleurosaur Neusticosaurus shows the plesiomorphic lamellar-zonal bone type and an incomplete endochondral ossification, like Nothosaurus. CONCLUSIONS/SIGNIFICANCE: Histological results hypothetically discussed in a phylogenetical context have a large impact on the current phylogenetic hypothesis of Sauropterygia, leaving the pachypleurosaurs polyphyletic. On the basis of histological data, Neusticosaurus would be related to Nothosaurus, whereas Anarosaurus would follow the pistosaur clade. Furthermore, the presence of fibrolamellar bone, which is accompanied with increased growth rates and presumably even with increased metabolic rates, already in Anarosaurus and Cymatosaurus can explain the success of the Pistosauroidea, the only sauropterygian group to survive into the Jurassic and give rise to the pelagic plesiosaur radiation.",
    url = "https://doi.org/10.1371/journal.pone.0011613",
    doi = "10.1371/journal.pone.0011613",
    openalex = "W2003799290",
    references = "doi1010079781489953919, doi101007978148995740520, doi101016jtree200508012, doi101029sc005p0175, doi10108010635150590950326, doi101371journalpone0007390, doi1016660094837320080340247ositlb20co2, doi1023071447735, doi10560219780801881206, openalexw2259112626"
}

BACKGROUND: Dysalotosaurus lettowvorbecki is a small ornithopod dinosaur known from thousands of bones and several ontogenetic stages. It was found in a single locality within the Tendaguru Formation of southeastern Tanzania, possibly representing a single herd. Dysalotosaurus provides an excellent case study for examining variation in bone microstructure and life history and helps to unravel the still mysterious growth pattern of small ornithopods. METHODOLOGY/PRINCIPAL FINDINGS: Five different skeletal elements were sampled, revealing microstructural variation between individuals, skeletal elements, cross sectional units, and ontogenetic stages. The bone wall consists of fibrolamellar bone with strong variability in vascularization and development of growth cycles. Larger bones with a high degree of utilization have high relative growth rates and seldom annuli/LAGs, whereas small and less intensively used bones have lower growth rates and a higher number of these resting lines. Due to the scarcity of annuli/LAGs, the reconstruction of the life history of Dysalotosaurus was carried out using regularly developed and alternating slow and fast growing zones. Dysalotosaurus was a precocial dinosaur, which experienced sexual maturity at ten years, had an indeterminate growth pattern, and maximum growth rates comparable to a large kangaroo. CONCLUSIONS/SIGNIFICANCE: The variation in the bone histology of Dysalotosaurus demonstrates the influence of size, utilization, and shape of bones on relative growth rates. Annuli/LAGs are not the only type of annual growth cycles that can be used to reconstruct the life history of fossil vertebrates, but the degree of development of these lines may be of importance for the reconstruction of paleobehavior. The regular development of annuli/LAGs in subadults and adults of large ornithopods therefore reflects higher seasonal stress due to higher food demands, migration, and altricial breeding behavior. Small ornithopods often lack regularly developed annuli/LAGs due to lower food demands, no need for migration, and precocial behavior.

@article{doi101371journalpone0029958,
    author = "Hübner, Tom",
    title = "Bone Histology in Dysalotosaurus lettowvorbecki (Ornithischia: Iguanodontia) – Variation, Growth, and Implications",
    year = "2012",
    journal = "PLoS ONE",
    abstract = "BACKGROUND: Dysalotosaurus lettowvorbecki is a small ornithopod dinosaur known from thousands of bones and several ontogenetic stages. It was found in a single locality within the Tendaguru Formation of southeastern Tanzania, possibly representing a single herd. Dysalotosaurus provides an excellent case study for examining variation in bone microstructure and life history and helps to unravel the still mysterious growth pattern of small ornithopods. METHODOLOGY/PRINCIPAL FINDINGS: Five different skeletal elements were sampled, revealing microstructural variation between individuals, skeletal elements, cross sectional units, and ontogenetic stages. The bone wall consists of fibrolamellar bone with strong variability in vascularization and development of growth cycles. Larger bones with a high degree of utilization have high relative growth rates and seldom annuli/LAGs, whereas small and less intensively used bones have lower growth rates and a higher number of these resting lines. Due to the scarcity of annuli/LAGs, the reconstruction of the life history of Dysalotosaurus was carried out using regularly developed and alternating slow and fast growing zones. Dysalotosaurus was a precocial dinosaur, which experienced sexual maturity at ten years, had an indeterminate growth pattern, and maximum growth rates comparable to a large kangaroo. CONCLUSIONS/SIGNIFICANCE: The variation in the bone histology of Dysalotosaurus demonstrates the influence of size, utilization, and shape of bones on relative growth rates. Annuli/LAGs are not the only type of annual growth cycles that can be used to reconstruct the life history of fossil vertebrates, but the degree of development of these lines may be of importance for the reconstruction of paleobehavior. The regular development of annuli/LAGs in subadults and adults of large ornithopods therefore reflects higher seasonal stress due to higher food demands, migration, and altricial breeding behavior. Small ornithopods often lack regularly developed annuli/LAGs due to lower food demands, no need for migration, and precocial behavior.",
    url = "https://doi.org/10.1371/journal.pone.0029958",
    doi = "10.1371/journal.pone.0029958",
    openalex = "W2088100200",
    references = "doi101016s1631069102014294, doi101017s0094837300021308, doi101029sc005p0175, doi10103835086558, doi101038nature02699, doi101038nature04633, doi101073pnas0708903105, doi101111j10963642201000620x, doi101111j146979981985tb04915x, doi101371journalpone0011613, doi1016660094837320000260466lhotts20co2, doi1016710272463420000200115lbhoth20co2"
}

@article{köhler2012seasonal,
    author = "Köhler, Meike and Marín-Moratalla, Nekane and Jordana, Xavier and Aanes, Ronny",
    title = "Seasonal bone growth and physiology in endotherms shed light on dinosaur physiology",
    year = "2012",
    journal = "Nature",
    url = "https://doi.org/10.1038/nature11264",
    doi = "10.1038/nature11264",
    number = "7407",
    openalex = "W2087820262",
    pages = "358-361",
    volume = "487",
    references = "doi101007s0044201016215, doi101016s1095643303003118, doi101016s1631069102014294, doi101017s1751731107000262, doi101111j13652435201001806x, doi101111j160005872000tb00300x, doi101643004585112002002117020co2, doi1016660094837320080340247ositlb20co2, doi1023071445584, doi105194hess1116332007"
}

The interest in mammalian palaeohistology has increased dramatically in the last two decades. Starting in 1849 via descriptive approaches, it has been demonstrated that bone tissue and vascularisation types correlate with several biological variables such as ontogenetic stage, growth rate, and ecology. Mammalian bone displays a large variety of bone tissues and vascularisation patterns reaching from lamellar or parallel-fibred to fibrolamellar or woven-fibred bone, depending on taxon and individual age. Here we systematically review the knowledge and methods on cynodont and mammalian bone microstructure as well as palaeohistology and discuss potential future research fields and techniques. We present new data on the bone microstructure of two extant marsupial species and of several extinct continental and island placental mammals. Extant marsupials display mainly parallel-fibred primary bone with radial and oblique but mainly longitudinal vascular canals. Three juvenile specimens of the dwarf island hippopotamid Hippopotamus minor from the Late Pleistocene of Cyprus show reticular to plexiform fibrolamellar bone. The island murid Mikrotia magna from the Late Miocene of Gargano, Italy displays parallel-fibred primary bone with reticular vascularisation and strong remodelling in the middle part of the cortex. Leithia sp., the dormouse from the Pleistocene of Sicily, is characterised by a primary bone cortex consisting of lamellar bone and a high amount of compact coarse cancellous bone. The bone cortex of the fossil continental lagomorph Prolagus oeningensis and three fossil species of insular Prolagus displays mainly parallel-fibred primary bone and reticular, radial as well as longitudinal vascularisation. Typical for large mammals, secondary bone in the giant rhinocerotoid Paraceratherium sp. from the Late Oligocene of Turkey is represented by dense Haversian bone. The skeletochronological features of Sinomegaceros yabei, a large-sized deer from the Pleistocene of Japan closely related to Megaloceros, indicate a high growth rate. These examples and the synthesis of existing data show the potential of bone microstructure to reveal essential information on life history evolution. The bone tissue and the skeletochronological data of the sampled island species suggest the presence of various modes of bone histological modification and mammalian life history evolution on islands to depend on factors of island evolution such as island size, distance from mainland, climate, phylogeny, and time of evolution.

@article{doi107717peerj1358,
    author = "Kolb, Christian and Scheyer, Torsten M. and Veitschegger, Kristof and Forasiepi, Analía M. and Amson, Eli and van der Geer, Alexandra and van den Hoek Ostende, Lars W. and Hayashi, Shoji and Sánchez‐Villagra, Marcelo R.",
    title = "Mammalian bone palaeohistology: a survey and new data with emphasis on island forms",
    year = "2015",
    journal = "PeerJ",
    abstract = "The interest in mammalian palaeohistology has increased dramatically in the last two decades. Starting in 1849 via descriptive approaches, it has been demonstrated that bone tissue and vascularisation types correlate with several biological variables such as ontogenetic stage, growth rate, and ecology. Mammalian bone displays a large variety of bone tissues and vascularisation patterns reaching from lamellar or parallel-fibred to fibrolamellar or woven-fibred bone, depending on taxon and individual age. Here we systematically review the knowledge and methods on cynodont and mammalian bone microstructure as well as palaeohistology and discuss potential future research fields and techniques. We present new data on the bone microstructure of two extant marsupial species and of several extinct continental and island placental mammals. Extant marsupials display mainly parallel-fibred primary bone with radial and oblique but mainly longitudinal vascular canals. Three juvenile specimens of the dwarf island hippopotamid Hippopotamus minor from the Late Pleistocene of Cyprus show reticular to plexiform fibrolamellar bone. The island murid Mikrotia magna from the Late Miocene of Gargano, Italy displays parallel-fibred primary bone with reticular vascularisation and strong remodelling in the middle part of the cortex. Leithia sp., the dormouse from the Pleistocene of Sicily, is characterised by a primary bone cortex consisting of lamellar bone and a high amount of compact coarse cancellous bone. The bone cortex of the fossil continental lagomorph Prolagus oeningensis and three fossil species of insular Prolagus displays mainly parallel-fibred primary bone and reticular, radial as well as longitudinal vascularisation. Typical for large mammals, secondary bone in the giant rhinocerotoid Paraceratherium sp. from the Late Oligocene of Turkey is represented by dense Haversian bone. The skeletochronological features of Sinomegaceros yabei, a large-sized deer from the Pleistocene of Japan closely related to Megaloceros, indicate a high growth rate. These examples and the synthesis of existing data show the potential of bone microstructure to reveal essential information on life history evolution. The bone tissue and the skeletochronological data of the sampled island species suggest the presence of various modes of bone histological modification and mammalian life history evolution on islands to depend on factors of island evolution such as island size, distance from mainland, climate, phylogeny, and time of evolution.",
    url = "https://doi.org/10.7717/peerj.1358",
    doi = "10.7717/peerj.1358",
    openalex = "W2109060835",
    references = "doi101002ajpa1330230215, doi1010079781489957405, doi101016s0379073800003765, doi101029sc005p0175, doi101093nargks1155, doi101126science1211028, doi101126science1229237, doi1023075403, doi107312kiel11918"
}

@article{doi101016jpalaeo201711041,
    author = "Kendall, Christopher and Eriksen, Anne and Kontopoulos, Ioannis and Collins, Matthew J. and Turner‐Walker, Gordon",
    title = "Diagenesis of archaeological bone and tooth",
    year = "2017",
    journal = "Palaeogeography Palaeoclimatology Palaeoecology",
    url = "https://doi.org/10.1016/j.palaeo.2017.11.041",
    doi = "10.1016/j.palaeo.2017.11.041",
    openalex = "W2769553357",
    references = "doi10100797836624715004, doi101016b9780125286282x50376, doi101016s0379073800003765, doi101017s0094837300005820, doi101021bi00769a018, doi101038171737a0, doi101038362709a0, doi101146annurevbiochem77032207120833, doi101146annurevbioeng8061505095721, doi101146annurevgenet37110801143214"
}