Abstract New discoveries on the ichnological site known as “the Pterosaur Beach of Crayssac” (lower Tithonian, Upper Jurassic; south-western France) answer the question of terrestrial capabilities of non-pterodactyloid pterosaurs. If the terrestrial type of locomotion of pterodactyloid pterosaurs has been solved from ichnological evidence for more than twenty years, no tracks and trackways referable to non-pterodactyloid pterosaurs have ever been described. Thus, the debate on terrestrial capabilities of these non-pterodactyloids was based on morpho-functional studies, with the main conclusion that those pterosaurs were arboreal dwellers and bad walkers. Six trackways referable to three non-pterodactyloid new ichnotaxa, maybe closely related to Rhamphorhynchidae, are described in this work. Their study leads to the conclusion that grounded non-pterodatyloids, at least during the Late Jurassic, were quadrupedal with digitigrade manus and plantigrade to digitigrade pes. They were clearly good walkers, even if hindlimbs are supposed to be hampered by the uropatagium, what could have constrained the terrestrial agility of these animals. Thus, from ichnological evidence and contrary to the current hypotheses, non-pterodactyloid pterosaurs seem to have been good walkers even though their trackways are very rare or unidentified to date. This rarity could be due to behaviour rather than to functional capacities, many non-pterodactyloids being considered both littoral fishers and arboreal or cliff dwellers. However, the concept of non-pterodactyloid “good climbers and bad walkers” has to be modified to “good climbers and rare walkers”, unless many non-pterodactyloid ichnites have yet to be discovered.

中文翻译：

---

首个非翼龙类翼龙类轨道和非翼龙类翼龙类的陆生能力

摘要 在被称为“克雷萨克翼龙海滩”（下提通阶，上侏罗纪；法国西南部）的考古遗址上的新发现回答了非翼龙类翼龙的陆地能力问题。如果从二十多年的时间证据中解决了翼手龙类翼龙的陆地运动类型，那么就没有描述过与非翼手龙类翼龙有关的轨迹和轨道。因此，关于这些非翼手龙的陆生能力的争论是基于形态功能研究的，主要结论是这些翼龙是树栖居民和不良步行者。在这项工作中描述了六个与三个非翼龙类新鱼纲有关的轨道，可能与 Rhamphorhynchidae 密切相关。他们的研究得出的结论是，至少在侏罗纪晚期，接地的非翼手龙是四足动物，手足类和足类为四足类。他们显然是很好的步行者，即使后肢应该受到 uropatagium 的阻碍，这可能会限制这些动物在陆地上的敏捷性。因此，从 ichnological 证据和目前的假设相反，非翼龙类翼龙似乎是很好的步行者，尽管它们的轨迹迄今为止非常罕见或未被识别。这种稀有性可能是由于行为而不是功能，许多非翼手龙类动物被认为是沿海渔民和树栖或悬崖居民。然而，非翼手类动物的“好登山者和坏步行者”的概念必须修改为“好登山者和稀有步行者”，