Hoffman D K, Goldsmith E R, Houssaye A, Maidment S C R, Felice R N, Mannion P D
Department of Earth Sciences, University College London, London, UK.
Department of Geosciences, Virginia Tech, Blacksburg, Virginia, USA.
J Anat. 2025 Jul;247(1):165-178. doi: 10.1111/joa.14231. Epub 2025 Feb 9.
Among living crocodylians, alligatoroids exhibit a wide range of body sizes and a biogeographic distribution that spans tropical-to-subtropical climates. The fossil record of alligatoroids, however, reveals even greater diversity, including multiple examples of gigantism and a broader distribution that extends into polar latitudes. Osteohistological studies on extant alligatoroids show that living alligators and caimans both exhibit seasonal growth, with roughly comparable growth rates. However, alligators and caimans diverged from one another over 60 million years ago; the dearth of studies on extinct alligatoroids makes it unclear if the shared condition in extant taxa reflects convergent responses to rapid climatic changes in the recent past or represents the ancestral condition in alligatoroids. Additionally, sample sizes are often limited to one or two individuals, especially in extinct crocodylians, obscuring any intraspecific variation present. To address this uncertainty, we conducted the largest monospecific osteohistological study of an extinct crocodylian to date, based on a sample of nine femora, providing unique insight into the intraspecific variation in growth of the early-diverging alligatoroid Diplocynodon hantoniensis from the late Eocene of the UK. The bone microanatomy of D. hantoniensis shows moderate compactness, with a well-defined medullary cavity, and osteohistological features that are generally consistent with those of extant alligatoroids. Samples vary greatly along a continuum in the degree of remodelling and vascularity, highlighting both the importance of evaluating intraspecific variation and limitations of basing histological assessments on singleton samples. Ontogenetic assessment indicates that our sample captures a range of skeletally immature to mature individuals, approximately corresponding to femoral size, but with notable exceptions possibly driven by sexual dimorphism. Body size estimates for D. hantoniensis (1.2-3.4 m) fall within the typical range of living American alligators (Alligator mississippiensis). Reconstruction from cyclical growth marks indicates a similar overall growth rate between D. hantoniensis and A. mississippiensis. As in extant alligatoroids more generally, this is determinate, seasonally-controlled growth. Femoral circumference scales positively with femoral length in D. hantoniensis, demonstrating similar allometry to A. mississippiensis. This differs from some other extant crocodylians (e.g. Crocodylus niloticus and Crocodylus johnstoni) and suggests conservation of allometric relationships in alligatoroids. This in-depth look into an early diverging alligatoroid indicates that seasonality and growth rates present in extant members were established near the base of the clade. Furthermore, it highlights the importance of including larger samples of singular species in order to capture potential variation when making clade-wide interpretations.
在现存的鳄形类动物中,短吻鳄类呈现出广泛的体型范围和跨越热带至亚热带气候的生物地理分布。然而,短吻鳄类的化石记录显示出更大的多样性,包括多个巨型化的例子以及延伸到极地纬度的更广泛分布。对现存短吻鳄类的骨组织学研究表明,现存的短吻鳄和凯门鳄都表现出季节性生长,生长速率大致相当。然而,短吻鳄和凯门鳄在6000多万年前就彼此分化;对已灭绝短吻鳄类的研究匮乏,使得目前尚不清楚现存分类群中这种共同的情况是反映了对近期快速气候变化的趋同反应,还是代表了短吻鳄类的原始状态。此外,样本量通常限于一两个个体,尤其是在已灭绝的鳄形类动物中,这掩盖了存在的任何种内变异。为了解决这种不确定性,我们基于九个股骨样本,对一种已灭绝的鳄形类动物进行了迄今为止最大规模的单物种骨组织学研究,为来自英国始新世晚期的早期分化短吻鳄类汉氏双齿鳄(Diplocynodon hantoniensis)的生长种内变异提供了独特见解。汉氏双齿鳄的骨微观解剖结构显示出适度的致密性,髓腔界限清晰,其骨组织学特征总体上与现存短吻鳄类一致。样本在重塑程度和血管化程度的连续体上差异很大,这既凸显了评估种内变异的重要性,也表明了基于单一样本进行组织学评估的局限性。个体发育评估表明,我们的样本涵盖了一系列骨骼未成熟到成熟的个体,大致与股骨大小相对应,但有明显例外,可能是由两性异形驱动的。对汉氏双齿鳄的体型估计(1.2 - 3.4米)落在现存美国短吻鳄(密西西比鳄,Alligator mississippiensis)的典型范围内。从周期性生长痕迹重建表明,汉氏双齿鳄和密西西比鳄的总体生长速率相似。与现存短吻鳄类更普遍的情况一样,这是一种确定性的、受季节控制的生长。在汉氏双齿鳄中,股骨周长与股骨长度呈正相关,显示出与密西西比鳄相似的异速生长关系。这与其他一些现存鳄形类动物(如尼罗鳄Crocodylus niloticus和强斯顿鳄Crocodylus johnstoni)不同,表明短吻鳄类中异速生长关系具有保守性。对早期分化的短吻鳄类的深入研究表明,现存成员中存在的季节性和生长速率在该类群的基部附近就已确立。此外,它强调了在进行全类群解释时纳入单个物种的更大样本以捕捉潜在变异的重要性。