Sander P M
Paläontologisches Institut und Museum, Universität Zürich, Switzerland.
Philos Trans R Soc Lond B Biol Sci. 1989 Nov 30;325(1230):561-666. doi: 10.1098/rstb.1989.0103.
The largest and most diverse collection of Pachypleurosauridae (Nothosauria, Reptilia) comes from Monte San Giorgio, Switzerland. Several hundred complete skeletons were collected from four distinct horizons of bituminous limestones and shales of Anisian-Ladinian boundary to early Ladinian age (Middle Triassic). Serpianosaurus mirigiolensis comes from the oldest strata, the Grenzbitumenzone Beds. The three younger strata, all in the Lower Meride Limestone, yield three species of Neusticosaurus. Neusticosaurus pusillus comes from the Cava Inferiore horizon, Neusticosaurus peyeri, new species, from the Cava Superiore horizon, and Neusticosaurus edwardsii, new combination, from the Alla Cascina horizon. Neusticosaurus pusillus is biostratigraphically important because it is one of the rare species reported from both the Germanic and the Alpine Triassic. Neusticosaurus pusillus and N. peyeri are small and very similar in their anatomy. Neusticosaurus species are easiest separated by their number of presacral vertebrae. Ornamentation of the bone surface is distinctive for all four pachypleurosaurids. Soft parts are rarely preserved, except for one partial squamation. The biological age of Neusticosaurus individuals can be determined by skeletochronology (aging by bone annuli). Small species of Neusticosaurus were sexually mature after three to four years and lived for six to nine years. Taphonomic analysis of the small species indicates attritional mortality and suggests weak bottom currents in the Monte San Giorgio basin during early Ladinian times. Morphometric comparison of all four pachypleurosaurids indicates that the changing vertebral numbers between species are largely due to a change in number of segments. All Monte San Giorgio pachypleurosaurids are sexually dimorphic in forelimb development. Sex x has poorly differentiated and relatively short humeri whereas sex y has well differentiated and relatively long humeri. The sexes are of about the same size and represented in roughly equal numbers. Identification of gender was not possible. Good growth series, especially of Neusticosaurus peyeri, from embryo to large adult permitted qualitative and quantitative study of ontogeny. The skull grows with negative allometry; the humerus grows isometrically or with positive allometry, depending on sex and species; the femur grows isometrically. The adult size range in N. peyeri is considerably larger than in modern reptiles. The Monte San Giorgio pachypleurosaurids are a monophyletic group. The phylogeny of this group is congruent with the stratigraphic distribution of its members.
肿肋龙科(幻龙目,爬行纲)数量最多、种类最为多样的化石收藏来自瑞士的圣乔治山。从早拉丁期(中三叠世)至安尼西阶-拉丁阶边界的沥青灰岩和页岩的四个不同层位采集到了数百具完整骨架。米氏塞尔皮亚诺龙来自最古老的地层,即 Grenzbitumenzone 层。另外三个较年轻的地层均位于下梅里德灰岩中,产出了三种纽氏肿肋龙。短小纽氏肿肋龙来自下卡瓦层位,佩氏纽氏肿肋龙(新物种)来自上卡瓦层位,爱德华兹纽氏肿肋龙(新组合)来自阿拉卡斯奇纳层位。短小纽氏肿肋龙在生物地层学上具有重要意义,因为它是在日耳曼和阿尔卑斯三叠纪均有报道的罕见物种之一。短小纽氏肿肋龙和佩氏纽氏肿肋龙体型较小且解剖结构非常相似。纽氏肿肋龙属的物种最容易通过其荐前椎骨的数量来区分。所有四种肿肋龙科动物的骨表面纹饰都很独特。除了一块部分保存的鳞片外,软组织很少被保存下来。纽氏肿肋龙个体的生物学年龄可以通过骨骼年轮学(通过骨骼年轮确定年龄)来确定。纽氏肿肋龙的小物种在三到四年后性成熟,寿命为六到九年。对小物种的埋藏学分析表明存在磨蚀性死亡,并表明在拉丁期早期圣乔治山盆地底部水流较弱。对所有四种肿肋龙科动物的形态测量比较表明,物种之间椎体数量的变化主要是由于节段数量的变化。圣乔治山所有的肿肋龙科动物在前肢发育上都存在两性异形。雄性的肱骨分化较差且相对较短,而雌性的肱骨分化良好且相对较长。两性体型大致相同,数量也大致相等。无法进行性别鉴定。良好的生长序列,尤其是佩氏纽氏肿肋龙从胚胎到大型成年个体的序列,使得对个体发育进行定性和定量研究成为可能。头骨以负异速生长;肱骨等速生长或正异速生长,这取决于性别和物种;股骨等速生长。佩氏纽氏肿肋龙的成年体型范围比现代爬行动物大得多。圣乔治山的肿肋龙科动物是一个单系类群。该类群的系统发育与其成员的地层分布一致。