Character-Taxon Matrix TNT File from A second peirosaurid crocodyliform from the Mid-Cretaceous Kem Kem Group of Morocco and the diversity of Gondwanan notosuchians outside South America

Notosuchians are an extinct clade of terrestrial crocodyliforms with a particularly rich record in the late Early to Late Cretaceous (approx. 130–66 Ma) of Gondwana. Although much of this diversity comes from South America, Africa and Indo-Madagascar have also yielded numerous notosuchian remains. Three notosuchian species are currently recognized from the early Late Cretaceous (approx. 100 Ma) Kem Kem Group of Morocco, including the peirosaurid <i>Hamadasuchus rebouli</i>. Here, we describe two new specimens that demonstrate the presence of at least a fourth notosuchian species in this fauna. <i>Antaeusuchus taouzensis</i> n. gen. n. sp. is incorporated into one of the largest notosuchian-focused character-taxon matrices yet to be compiled, comprising 443 characters scored for 63 notosuchian species, with an increased sampling of African and peirosaurid species. Parsimony analyses run under equal and extended implied weighting consistently recover <i>Antaeusuchus</i> as a peirosaurid notosuchian, supported by the presence of two distinct waves on the dorsal dentary surface, a surangular which laterally overlaps the dentary above the mandibular fenestra, and a relatively broad mandibular symphysis. Within Peirosauridae, <i>Antaeusuchus</i> is recovered as the sister taxon of <i>Hamadasuchus</i>. However, it differs from <i>Hamadasuchus</i> with respect to several features, including the ornamentation of the lateral surface of the mandible, the angle of divergence of the mandibular rami, the texture of tooth enamel and the shape of the teeth, supporting their generic distinction. We present a critical reappraisal of the non-South American Gondwanan notosuchian record, which spans the Middle Jurassic–Late Eocene. This review, as well as our phylogenetic analyses, indicate the existence of at least three approximately contemporaneous peirosaurid lineages within the Kem Kem Group, alongside other notosuchians, and support the peirosaurid affinities of the ‘trematochampsid’ <i>Miadanasuchus oblita</i> from the Maastrichtian of Madagascar. Furthermore, the Cretaceous record demonstrates the presence of multiple lineages of approximately contemporaneous notosuchians in several African and Madagascan faunas, and supports previous suggestions regarding an undocumented pre-Aptian radiation of Notosuchia. By contrast, the post-Cretaceous record is depauperate, comprising rare occurrences of sebecosuchians in north Africa prior to their extirpation.

诺托鳄类（Notosuchians）是一类已灭绝的陆生鳄形类（terrestrial crocodyliforms）演化支，其化石记录在冈瓦纳大陆（Gondwana）的早白垩世晚期至晚白垩世（late Early to Late Cretaceous，约1.3亿至6600万年前）尤为丰富。尽管该类群的多数多样性化石产自南美洲，但非洲与印度-马达加斯加地块也产出了大量诺托鳄类化石遗存。目前在摩洛哥晚白垩世早期（约1亿年前）的凯姆凯姆组（Kem Kem Group）中已确认3种诺托鳄类，其中包括佩罗鳄科（peirosaurid）的<i>Hamadasuchus rebouli</i>。本文记述了两件新标本，证实该动物群中至少存在第四种诺托鳄类。新属新种<i>Antaeusuchus taouzensis</i> n. gen. n. sp.被纳入迄今规模最大的以诺托鳄类为核心的性状-分类矩阵（character-taxon matrix）之一，该矩阵包含针对63种诺托鳄类的443个性状记录，并增加了非洲类群与佩罗鳄科物种的采样量。在等权与扩展隐含加权（equal and extended implied weighting）条件下运行的简约分析（parsimony analyses）均将<i>Antaeusuchus</i>恢复为佩罗鳄科诺托鳄类，这一结果得到多项特征的支持：齿骨背侧表面存在两处独立的隆起，下颌孔上方的上隅骨侧向覆盖齿骨，以及相对宽阔的下颌联合（mandibular symphysis）。在佩罗鳄科内部，<i>Antaeusuchus</i>与<i>Hamadasuchus</i>互为姊妹群（sister taxon）。但<i>Antaeusuchus</i>与<i>Hamadasuchus</i>在多项特征上存在差异，包括下颌侧面的纹饰、下颌支（mandibular rami）的发散角度、牙釉质纹理以及牙齿形态，这些差异支持二者分属不同的属。本文对冈瓦纳大陆非南美洲的诺托鳄类化石记录进行了批判性重新评述，该记录的时代跨度从中侏罗世至晚始新世。本次评述与系统发育分析均表明，凯姆凯姆组中至少存在3个近同期的佩罗鳄科演化支，伴生有其他诺托鳄类；同时支持产自马达加斯加马斯特里赫特阶（Maastrichtian）的“颤鳄科”（trematochampsid）<i>Miadanasuchus oblita</i>属于佩罗鳄科。此外，白垩纪的化石记录显示，非洲与马达加斯加的多个动物群中存在多个近同期的诺托鳄类演化支，并支持此前提出的诺托鳄类在阿普第阶（Aptian）之前存在未被记录的辐射演化事件。与之形成对比的是，晚白垩世后的化石记录极为贫乏，仅在北非存在少量西贝鳄类（sebecosuchians）的记录，这类群最终在此区域局部灭绝。