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Fossil moray eels (Muraenidae) from the interoceanic Central American seaway 中美洲海道的海鳗化石(海鳗科)
IF 1.6 4区 地球科学 Q2 PALEONTOLOGY Pub Date : 2025-08-01 Epub Date: 2025-05-22 DOI: 10.1016/j.geobios.2024.12.004
Orangel Aguilera , Carlos de Gracia , Félix Rodriguez , Olga Oliveira de Araújo , Werner Schwarzhans , Philippe Béarez , Antoni Lombarte , Paulo Andreas Buckup , Ricardo Tadeu Lopes
Despite the wide distribution and diversity of extant Anguilliformes in tropical America (central western Atlantic and central eastern Pacific), no fossil moray ell (Muraenidae) has been reported to date from this region. Here, we describe the first three fossil specimens of Muraeninae from America: one from the Late Miocene Gatun Formation (Fm.), the second from the Late Pliocene Escudo de Veraguas Fm., both from Panama, and the third from the Late Pliocene Rio Banano Fm. in Costa Rica. Gymnothorax pierreolivieri nov. sp. from the Gatun Fm. is described based on the skull and an in situ otolith. The new species is characterized by a premaxilla-ethmo-vomerine bone (PMx-Etv) with 15 marginal fang-like teeth, two medial teeth, and 13 vomerine teeth arranged in a single row. The dorsal stem of the PMx-Etv forms a very thick and massive structure, with a single anterior foramen located in the anterior tip of the bone. The maxilla has 16 teeth arranged in a double row. The sagitta otolith is elliptic and tapered anteriorly; its posterior margin is smooth and gently rounded; its dorsal margin is slightly arched and elevated posteriorly. The isolated otoliths from the Escudo de Veraguas Fm. in Panama and from the Rio Banano Fm. in Costa Rica are putatively assigned to Echidna sp. and Enchelycore sp., respectively. The diversification of American Muraenidae seems to have been driven by oceanic water interchanges and species dispersal associated to marine currents that flowed eastward through the Central American Seaway prior to the total closure of the Panama Isthmus. The extant American muraenid diversity illustrates allopatric speciation (vicariance) when populations from the Atlantic and the Pacific oceans were isolated by the late Tertiary uplift of the Isthmus of Panama. The palaeoceanographic changes driven by this geological event have had evolutionary consequences on faunal turnover and extinctions through space and time and is reflected by the extant fish diversity.
尽管在热带美洲(大西洋中西部和太平洋中部东部)现存的鳗形目分布广泛,种类繁多,但迄今为止在该地区尚未发现鳗形目(Muraenidae)化石。在这里,我们描述了来自美洲的前三个Muraeninae化石标本:一个来自晚中新世的Gatun组(Fm.),另一个来自晚上新世的Escudo de Veraguas组。第三个来自晚上新世的Banano Fm。在哥斯达黎加。裸子胸(Gymnothorax pierreolivieri) 11月11日摄于加顿Fm。是根据颅骨和原位耳石描述的。新物种的特征是前颌-下颌-下颌骨(PMx-Etv), 15个边缘尖牙,2个中间齿和13个排成一行的下颌齿。PMx-Etv的背干形成一个非常厚且巨大的结构,在骨的前尖端有一个单一的前孔。上颌骨有16颗牙齿排成两排。矢状耳石呈椭圆形,前呈锥形;它的后缘光滑,微圆;它的背缘稍呈弓形并向后抬高。从Escudo de Veraguas Fm发现的孤立耳石。在巴拿马和bbbbo Banano Fm。在哥斯达黎加分别被推定为Echidna sp和Enchelycore sp。美洲Muraenidae的多样化似乎是由海水交换和物种分散所驱动的,在巴拿马地峡完全关闭之前,洋流向东流过中美洲海道。现存的美洲muraenid多样性说明了来自大西洋和太平洋的种群被晚第三纪巴拿马地峡的隆起所隔离时的异域物种形成(vicariance)。由这一地质事件驱动的古海洋学变化,在空间和时间上对动物更替和灭绝产生了进化后果,并反映在现存鱼类多样性上。
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引用次数: 0
Borophagine canids of the Monarch Mill Formation (Middle Miocene), Nevada, U.S.A. 美国内华达帝王磨坊组(中新世中期)的犬科动物
IF 1.6 4区 地球科学 Q2 PALEONTOLOGY Pub Date : 2025-08-01 Epub Date: 2025-05-22 DOI: 10.1016/j.geobios.2024.11.011
Z. Jack Tseng , Xiaoming Wang
The non-marine Middle Miocene Monarch Mill Formation (Nevada, U.S.A.) outcrops in the Middlegate Basin and preserves fossil mammals that lived during the establishment of the present-day basin and range topography in central Nevada. The Monarch Mill Formation and its mammal fauna overlie the Middlegate Formation and its flora, and together the biota provides an informative window into ecosystem composition at the geographic crossroads between the Great Plains and the coastal regions of North America during the Middle Miocene. Here we report on previously undescribed and/or unfigured canids from the Monarch Mill Formation. To the previously known borophagine Tomarctus brevirostris and the fox-like canine Leptocyon we add two borophagine genera to the Eastgate Local Fauna of the Monarch Mill Formation. Specimens of the hypocarnivores Paracynarctus and Cynarctus are described and provide an enhanced understanding of the Middlegate Basin canid assemblage, now comprising at least four genera. Together with floral evidence, this canid assemblage is indicative of mesocarnivore-sustaining vegetation and locally limited open environments, with no hypercarnivorous canid occurrences. Rapid regional subsidence was an overarching factor in the shift towards present-day hyper- and meso-carnivore dominated canid assemblages in the Basin and Range Province, and the appearance of hypercarnivores in post-Barstovian times may reflect contemporaneous regional topographic shifts in the Great Basin at large.
非海相中中新世帝王磨坊组(美国内华达州)在米德尔盖特盆地露头,并保存了在内华达中部盆地和山脉地形建立期间生活的哺乳动物化石。君主磨坊组和它的哺乳动物群覆盖在米德尔盖特组和它的植物群上,这些生物群一起提供了一个信息窗口,可以了解中新世中期大平原和北美沿海地区地理十字路口的生态系统组成。在这里,我们报告了以前未描述和/或未计算的犬科动物从君主磨坊组。在先前已知的borophagine Tomarctus brevirostris和fox-like canine lepptocyon的基础上,我们将两个borophagine属添加到Monarch Mill Formation的Eastgate Local Fauna中。描述了准食肉动物Paracynarctus和Cynarctus的标本,并提供了对米德尔盖特盆地犬科动物组合的更好理解,现在至少包括四个属。结合植物的证据,这一犬科动物组合表明了中肉食性植被和局部有限的开放环境,没有超肉食性犬科动物的出现。快速的区域沉降是盆地和山脉省向现今超食肉动物和中食肉动物为主的犬科动物组合转变的主要因素,后巴斯托夫时代超食肉动物的出现可能反映了大盆地同期区域地形的变化。
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引用次数: 0
Diversity of the pterodactyloid ichnites of Crayssac (Lower Tithonian, Late Jurassic, southwestern France) 克雷萨克(法国西南部晚侏罗世下梯东世)翼手类岩的多样性
IF 1.6 4区 地球科学 Q2 PALEONTOLOGY Pub Date : 2025-08-01 Epub Date: 2025-05-22 DOI: 10.1016/j.geobios.2024.12.002
Jean-Michel Mazin, Joane Pouech
The ichnosite of Crayssac (Upper Jurassic, southwestern France) yields numerous vertebrates and invertebrates trackways, among which the pterodactyloid pterosaur trackways are frequent. One hundred and sixty nine vertebrate trackways (turtles, crocodilians, dinosaurs and pterosaurs) have been unearthed by our team. They are preserved in situ, are sheltered by a building and legally protected within the Réserve Naturelle Nationale Géologique du Lot. Among these vertebrate trackways, sixty-four can be attributed to tiny- to medium-sized pterodactyloids, representing five morphotypes referred to five new ichnospecies: Morphotype I, Pteraichnus occitanis (1 specimen); Morphotype II, Pteraichnus cadurcii (12 specimens); Morphotype III, Pteraichnus communis (21 specimens); Morphotype IV, Pteraichnus dichnopollex (10 specimens); and Morphotype V, Pteraichnus pyrenaicus (7 specimens). These trackways are finely preserved and represent a unique, abundant and diversified sample. Study of complete trackways rather than isolated footprints allows access to important parameters such as the velocity or the comparative gauges. Similarly, the orientation of all vertebrate trackways shows that the pterosaurs moved actively across the mudflat, without preferred direction.
Crayssac(法国西南部上侏罗统)的ichnosite上发现了许多脊椎动物和无脊椎动物的足迹,其中翼手类翼龙足迹较为常见。我们的团队已经发现了169条脊椎动物的足迹(海龟、鳄鱼、恐龙和翼龙)。它们被保存在原地,受到建筑的保护,并受到国家自然保护组织(r serve Naturelle Nationale g ologique du Lot)的法律保护。在这些脊椎动物足迹中,64条属于中小型翼手纲,代表5种形态类型,涉及5个新种:形态类型I, Pteraichnus occitanis(1个标本);形态型II型,无迹蝶(12份);3型,共翼毛虫(21份);形态型IV,双斑翼蚊(10份);V型为pyrenaichnus(7份)。这些铁轨保存完好,代表了一个独特、丰富和多样化的样本。研究完整的轨道,而不是孤立的足迹,可以获得重要的参数,如速度或比较仪表。同样,所有脊椎动物足迹的方向都表明翼龙在泥滩上主动移动,没有偏好的方向。
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引用次数: 0
Doing the time warp again: Electron Spin Resonance dating reveals oldest numeric age for Notiomastodon platensis Ameghino, 1888 (Mammalia, Proboscidea) 再次进行时间扭曲:电子自旋共振测年揭示了1888年Notiomastodon platensis Ameghino最古老的数字年龄(哺乳动物,长鼻目)
IF 1.6 4区 地球科学 Q2 PALEONTOLOGY Pub Date : 2025-08-01 Epub Date: 2025-05-22 DOI: 10.1016/j.geobios.2024.11.008
Dimila Mothé , Angela Kinoshita , Oswaldo Baffa , Carlos A. Luna
Here we used Electron Spin Resonance (ESR) dating methods on seven fossil specimens to update the temporal and geographic distributions of the Quaternary proboscidean Notiomastodon platensis Ameghino 1888, from Córdoba Province, Argentina. While abundant in the Late Pleistocene/Early Holocene South American fossil record, the knowledge about the Early-Middle Pleistocene records of this proboscidean is scarce due to limited numeric datings data. ESR results reveal numeric ages ranging from 560 ± 40 to 47 ± 7 ka, placing the species within the Ensenadan to Lujanian stages of the Pleistocene (Chibanian to Late Pleistocene). The Ensenadan record represents the oldest numeric age of Notiomastodon platensis in South America. The study highlights the importance of numeric dating in addressing the geochronological data gap for South American megafauna and reveals the multiple environments that Notiomastodon platensis inhabited during Quaternary, suggesting slow vertical migrations in response to climatic changes, with mountainous regions of Cordoba province serving as refuges. The need for further numeric datings is emphasized in this study, to improve our understanding of the evolutionary history and extinction drivers of South American proboscideans during the Quaternary.
本文利用电子自旋共振(ESR)测年方法,对阿根廷Córdoba省第四纪长鼻目Notiomastodon platensis Ameghino 1888化石标本进行了时间和地理分布的更新。虽然南美晚更新世/全新世早期化石记录丰富,但由于数值测年数据有限,对该类早-中更新世化石记录的了解很少。ESR结果显示,该物种的年龄范围为560±40 ~ 47±7 ka,处于更新世的Ensenadan至Lujanian阶段(Chibanian至晚更新世)。Ensenadan记录代表了南美洲Notiomastodon platensis最古老的数字年龄。该研究强调了数值测年在解决南美巨型动物地质年代学数据缺口方面的重要性,揭示了第四纪中非乳齿象(Notiomastodon platensis)居住的多种环境,表明它们在气候变化的影响下进行了缓慢的垂直迁移,科尔多瓦省的山区是它们的避难所。本研究强调需要进一步的数值定年,以提高我们对第四纪南美长鼻目动物的进化史和灭绝驱动因素的理解。
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引用次数: 0
Mississippian-Lower Pennsylvanian foraminifers from the Sierra Nacimiento and San Pedro Mountains, North-Central New Mexico, USA 来自美国新墨西哥州中北部的Sierra Nacimiento和San Pedro山脉的密西西比-下宾夕法尼亚有孔虫
IF 1.6 4区 地球科学 Q2 PALEONTOLOGY Pub Date : 2025-08-01 Epub Date: 2025-05-22 DOI: 10.1016/j.geobios.2024.11.009
Karl Krainer , Spencer G. Lucas , Daniel Vachard
In the Sierra Nacimiento-San Pedro Mountains-Jemez Mountains of north-central New Mexico (USA), the thin succession of Mississippian sediments is termed the Arroyo Peñasco Formation, overlain by the Log Springs Formation. The Arroyo Peñasco Formation is subdivided into Del Padre Member composed of siliciclastic sediments and the overlying Espiritu Santo Member composed of carbonate sediments. Foraminifers of the Espiritu Santo Member at Lion Spring in the northern San Pedro Mountains indicate a Tournaisian age. The lower Tournaisian (= Hastarian = biozones MFZ1-4 of Poty et al., 2006 [Geological Magazine 143, 829–857]) is locally characterized by Septaglomospiranella ex gr. primaeva, Septabrunsiina minuta and Tournayella sp. The upper Tournaisian (= Ivorian = biozones MFZ5-9 of Poty et al., 2006) is characterized by Spinoendothyra, Inflatoendothyra and Tuberendothyra. In the Sierra Nacimiento-San Pedro Mountains-Jemez Mountains, the sediments of the Arroyo Peñasco Group are unconformably overlain by nonmarine redbeds of the Log Springs Formation (Serpukhovian/upper Chesterian). Some of the carbonate clasts of the basal conglomerate of the Log Springs Formation at Lion Spring contain late early Viséan (biozone MFZ11B of Poty et al., 2006) foraminifers and algae, including Koninckopora and Paraarchaediscus. These carbonate clasts are reworked from the underlying Arroyo Peñasco Formation (Espiritu Santo Member). In north-central New Mexico, a transgression during the Early Pennsylvanian (Morrowan) caused deposition of the shallow marine sediments of the Osha Canyon Formation. The Morrowan age of the Osha Canyon Formation is based on Nigrispiroides nov. gen., Iriclinella, Globivalvulina, and Millerella.
在美国新墨西哥州中北部的Sierra Nacimiento-San Pedro Mountains- jemez Mountains,密西西比沉积的薄层序列被称为Arroyo Peñasco组,其上覆盖着Log Springs组。Arroyo Peñasco组可分为Del Padre组和espiiritu Santo组,其上为碳酸盐沉积。圣佩德罗山脉北部狮子泉的埃斯皮里图圣成员的有孔虫表明了图尔纳时代。图尔奈世下部(= Hastarian = MFZ1-4, Poty et al, 2006)局部特征为Septaglomospiranella ex gr. primaeva、Septabrunsiina minuta和Tournayella sp.;图尔奈世上部(= Ivorian = MFZ5-9, Poty et al, 2006)局部特征为Spinoendothyra、Inflatoendothyra和Tuberendothyra。在Sierra Nacimiento-San Pedro Mountains- jemez Mountains, Arroyo Peñasco群的沉积物被Log Springs组(Serpukhovian/upper Chesterian)的非海相红层不整合覆盖。狮子泉Log Springs组基底砾岩的部分碳酸盐岩碎屑中含有晚、早vis (Poty et al., 2006的MFZ11B生物带)有孔虫和藻类,包括Koninckopora和Paraarchaediscus。这些碳酸盐碎屑来自下部的阿罗约Peñasco组(圣埃斯皮里图成员)。在新墨西哥州中北部,早宾夕法尼亚(Morrowan)时期的一次海侵导致了奥沙峡谷组浅海沉积物的沉积。Osha峡谷组的Morrowan时代是基于Nigrispiroides nov. gen.、Iriclinella、Globivalvulina和Millerella。
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引用次数: 0
Megaglomerospora lealiae nov. gen., nov. sp. from the upper Carboniferous of Portugal: the largest glomeromycotan fungal spores 来自葡萄牙上石炭世的叶状巨聚孢:最大的肾小球真菌孢子
IF 1.6 4区 地球科学 Q2 PALEONTOLOGY Pub Date : 2025-08-01 Epub Date: 2025-05-22 DOI: 10.1016/j.geobios.2024.11.010
Pedro Correia , Artur A. Sá , Zélia Pereira
A new genus and species of fossil fungus, Megaglomerospora lealiae, is described from the Buçaco Carboniferous Basin (upper Stephanian C, Upper Pennsylvanian, upper Carboniferous), in central western Portugal. The new fossil fungus consists of a dense cluster of silicified large spores. These new fungal spores are oblong, subelliptical to subspherical-shaped, with a glabrous surface characterized by having a lipid-filled lumen, and display a strong septate-like hypha attached. The presence of lobe-shaped germination shields suggests close affinities to Diversisporales (Glomeromycota). Megaglomerospora lealiae nov. gen., nov. sp. is remarkably distinctive because it is by far the largest fossil fungal spore (∼1.6 mm long) documented for the phylum Glomeromycota. This is the first report of an endomycorrhizal‐like fungus from the Carboniferous of Iberia.
在葡萄牙中西部的buaco石炭世盆地(上斯蒂芬纪C,上宾夕法尼亚纪,上石炭世)描述了一种新的真菌化石Megaglomerospora lealiae属。新的真菌化石由密集的硅化大孢子簇组成。这些新的真菌孢子呈椭圆形,近椭圆形至近球形,表面无毛,具有充满脂质的腔,并显示有很强的分离样菌丝附着。叶状萌发屏障的存在表明其与Diversisporales (Glomeromycota)有密切的亲缘关系。叶状巨聚孢菌(Megaglomerospora lealiae, 11 . gen., 11 . sp.)非常独特,因为它是迄今为止记录在案的肾小球菌门中最大的真菌孢子化石(约1.6毫米长)。这是来自伊比利亚石炭纪的一种类似内生菌根的真菌的首次报道。
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引用次数: 0
IF 1.6 4区 地球科学 Q2 PALEONTOLOGY Pub Date : 2025-06-12 DOI: 10.1016/j.geobios.2025.05.007
Mickaël Lhéritier
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引用次数: 0
IF 1.6 4区 地球科学 Q2 PALEONTOLOGY Pub Date : 2025-06-03 DOI: 10.1016/j.geobios.2025.05.001
Adolfo Pacheco-Castro
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引用次数: 0
Osteohistology of enantiornithine birds from the Lower Cretaceous Xiagou Formation 下白垩统下沟组反鸟目鸟类的骨组织学
IF 1.6 4区 地球科学 Q2 PALEONTOLOGY Pub Date : 2025-06-01 Epub Date: 2025-03-05 DOI: 10.1016/j.geobios.2024.08.020
Jessie Atterholt , Jingmai K. O’Connor , Hailu You
We describe the osteohistology of five enantiornithine bird specimens from Lower Cretaceous Xiagou Formation deposits of the Changma locality in northwestern Gansu Province, China. Samples were taken from the femora of: three specimens of Avimaia schweitzerae (IVPP V25371, IVPP V31956, and GSGM-04-CM-007), Qiliania graffini GSGM-04-CM-006, and Novavis pubisculata IVPP V31957. The objective of this study is to describe intrageneric variation (in Avimaia), and intertaxonomic variation among enantiornithine birds coexisting in an ecosystem. All five specimens have a femoral cortex composed mainly of parallel fibered bone with relatively low vascularity. All three Avimaia specimens have 2–3 vascular canals, and asymmetrical growth marks, indicating cortical drift. In Qiliania there are eight longitudinal vascular canals, five of which are concentrated in one region of the cortex. Although the gross anatomy of the skeleton and fusion of compound elements indicates morphological maturity, neither growth marks, an outer circumferential layer (OCL), nor an inner circumferential layer (ICL) are present. The femur of Novavis has some regions of a woven parallel complex and a higher level of vascularity relative to the other specimens (14 longitudinal channels present). Although this specimen is morphologically immature based on gross anatomy, the femur has a well-developed OCL and ICL. These results emphasize the enantiornithine offset between morphological maturity and osteohistological maturity. Development of the OCL appears to be decoupled from morphological maturity, in some cases forming before the skeleton has fully fused, and in others well after. The specimens are similar in size but vary considerably in the number of growth marks present, from none to two. This suggests either developmental plasticity and diverse growth strategies and, complicates attempts to interpret relative age and growth stage in enantiornithines.
本文描述了甘肃长马地区下白垩统下沟组5个反鸟目鸟类标本的骨组织学特征。采集的标本分别为:3只豚鼠(Avimaia schweitzerae) (IVPP V25371、IVPP V31956和GSGM-04-CM-007)、棘蝇(Qiliania graffini) (GSGM-04-CM-006)和诺维斯(Novavis pubisculata) (IVPP V31957)。本研究的目的是描述在一个生态系统中共存的反鸟目鸟类的属内变异(在Avimaia)和分类间变异。所有五个标本的股骨皮质主要由平行纤维骨组成,血管密度相对较低。三个Avimaia标本均有2-3条血管管,且有不对称的生长标记,表明皮层漂移。祁连菌有8条纵向维管,其中5条集中在皮层的一个区域。虽然骨骼的大体解剖和复合元素的融合表明形态成熟,但既没有生长标志,也没有外周层(OCL),也没有内周层(ICL)。与其他标本相比,Novavis的股骨具有编织平行复合体的一些区域和更高水平的血管性(存在14条纵向通道)。虽然该标本在大体解剖上形态不成熟,但股骨具有发育良好的OCL和ICL。这些结果强调形态成熟和骨组织成熟之间的反鸟氨酸偏移。OCL的发育似乎与形态成熟脱钩,在某些情况下,在骨骼完全融合之前形成,而在其他情况下,则在骨骼完全融合之后形成。这些标本在大小上相似,但在生长标记的数量上差别很大,从没有到两个不等。这表明发育可塑性和多样化的生长策略,并使解释对鸟的相对年龄和生长阶段的尝试复杂化。
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引用次数: 0
Craniocervical morphological integration in birds 鸟类颅颈形态整合
IF 1.6 4区 地球科学 Q2 PALEONTOLOGY Pub Date : 2025-06-01 Epub Date: 2025-03-26 DOI: 10.1016/j.geobios.2024.11.007
Jesús Marugán-Lobón , Sergio M. Nebreda
This study investigates the morphological diversity of the neck and its integration with skull anatomy in crown-group birds (Neornithes). The craniocervical articulation, where the skull connects with the neck, can adopt either caudal (rearward) or ventral (downward) orientations. Morphological coherence is crucial for maintaining head stability and precise movement, suggesting that the evolution of these structures must be not only functionally, but also developmentally coordinated (i.e., morphologically integrated). To explore this hypothesis, the relationship between conventional morphometric data of the neck and geometric morphometric data of the skull were assessed using multivariate statistics (Regressions and Two-block Partial Least Squares) across a broad phylogenetic range of Neornithes. Results indicate a significant level of integration between neck and skull morphologies, where variation in neck length, vertebral counts and relative lengths correspond predictably with specific craniocervical articulations. The most pronounced skull variation occurs around the occipital region, which we interpret as possibly relating to the mesodermic origin and shared genetic signalling in the morphogenesis of all craniocervical bones. Additionally, craniofacial changes align with craniocervical modifications, implying that the skull and neck evolve as a unified yet modular system. Analysis of selected fossils (Tsaagan, Archaeopteryx, and Pengornis) suggests that the ancestral craniocervical configuration was caudal, with neck changes associated with the development of a beak and craniocervical ventralization emerging in the lineage leading to modern birds.
本研究探讨了冠类鸟类颈部的形态多样性及其与颅骨解剖的结合。颅颈关节,即颅骨与颈部相连的部位,可采用尾侧(后)或腹侧(下)方向。形态一致性对于维持头部稳定性和精确运动至关重要,这表明这些结构的进化不仅必须在功能上协调,而且必须在发育上协调(即形态上的整合)。为了探索这一假设,我们使用多元统计(回归和双块偏最小二乘法)评估了颈部的常规形态测量数据和颅骨的几何形态测量数据之间的关系。结果表明,颈部和颅骨形态之间存在显著的整合水平,其中颈部长度、椎体计数和相对长度的变化可预测地与特定的颅颈关节相对应。最明显的颅骨变异发生在枕区周围,我们认为这可能与所有颅颈骨的中胚层起源和形态发生中共享的遗传信号有关。此外,颅面变化与颅颈变化一致,这意味着颅骨和颈部作为一个统一的模块化系统进化。对选定化石(Tsaagan,始祖鸟和Pengornis)的分析表明,祖先的头颈结构是尾端的,颈部的变化与喙的发育和头颈腹侧化有关,在导致现代鸟类的谱系中出现。
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