Abstract How species diversification occurs remains an unanswered question in predatory marine invertebrates, such as sea snails of the family Terebridae. However, the anatomical disparity found throughput the Terebridae provides a unique perspective for investigating diversification patterns in venomous predators. In this study, a new dated molecular phylogeny of the Terebridae is used as a framework for investigating diversification of the family through time, and for testing the putative role of intrinsic and extrinsic traits, such as shell size, larval ecology, bathymetric distribution, and anatomical features of the venom apparatus, as drivers of terebrid species diversification. Macroevolutionary analysis revealed that when diversification rates do not vary across Terebridae clades, the whole family has been increasing its global diversification rate since 25 Ma. We recovered evidence for a concurrent increase in diversification of depth ranges, while shell size appeared to have undergone a fast divergence early in terebrid evolutionary history. Our data also confirm that planktotrophy is the ancestral larval ecology in terebrids, and evolutionary modeling highlighted that shell size is linked to larval ecology of the Terebridae, with species with long-living pelagic larvae tending to be larger and have a broader size range than lecithotrophic species. Although we recovered patterns of size and depth trait diversification through time and across clades, the presence or absence of a venom gland (VG) did not appear to have impacted Terebridae diversification. Terebrids have lost their venom apparatus several times and we confirm that the loss of a VG happened in phylogenetically clustered terminal taxa and that reversal is extremely unlikely. Our findings suggest that environmental factors, and not venom, have had more influence on terebrid evolution.

中文翻译：

---

宏观进化分析表明，环境因素，而不是毒液装置，在陆生蜗牛的多样化中发挥着关键作用

摘要 对于捕食性海洋无脊椎动物，例如特雷布达科的海蜗牛，物种多样化是如何发生的仍然是一个悬而未决的问题。然而，截肢科动物的解剖学差异为研究有毒捕食者的多样化模式提供了独特的视角。在这项研究中，新的已过时的拟肢动物分子系统发育学被用作研究该家族随时间的多样化的框架，并测试内在和外在特征的假定作用，例如壳大小、幼虫生态学、水深分布和毒液装置的解剖学特征，作为三足动物物种多样化的驱动因素。宏观进化分析表明，当不同步履虫科进化枝的多样化率没有变化时，自 25 Ma 以来，整个家族的总体多样化率一直在增加。我们发现了深度范围多样化同时增加的证据，而贝壳尺寸似乎在三文鱼进化史的早期经历了快速分化。我们的数据还证实，浮游营养型是陆栖动物的祖先幼虫生态，进化模型强调，壳的大小与陆栖动物的幼虫生态有关，具有长寿的中上层幼虫的物种往往比卵食性幼虫更大，尺寸范围更广。物种。尽管我们恢复了随时间和跨分支的大小和深度性状多样化的模式，但毒液腺（VG）的存在或不存在似乎并没有影响Tebridae的多样化。Terebrids 已经多次失去毒液装置，我们证实 VG 的丢失发生在系统发育聚类的末端类群中，并且逆转的可能性极小。我们的研究结果表明，环境因素，而不是毒液，对三文鱼进化的影响更大。