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The First Reconstruction of the Head Anatomy of a Cretaceous Insect, †Gerontoformica gracilis (Hymenoptera: Formicidae), and the Early Evolution of Ants 白垩纪昆虫†Gerontoformica gracilis(膜翅目:蚁科)头部解剖结构的首次重建与蚂蚁的早期进化
IF 3.4 1区 农林科学 Pub Date : 2022-09-01 DOI: 10.1093/isd/ixac013
A. Richter, B. Boudinot, Shûhei Yamamoto, Julian Katzke, R. Beutel
Abstract The fossil record allows a unique glimpse into the evolutionary history of organisms living on Earth today. We discovered a specimen of the stem group ant †Gerontoformica gracilis (Barden and Grimaldi, 2014) in Kachin amber with near-complete preservation of internal head structures, which we document employing µ-computed-tomography-based 3D reconstructions. We compare †Gerontoformica to four outgroup taxa and four extant ant species, employing parsimony and Bayesian ancestral state reconstruction to identify morphological differences and similarities between stem and crown ants and thus improve our understanding of ant evolution through the lens of head anatomy. Of 149 morphological characters, 87 are new in this study, and almost all applicable to the fossil. †Gerontoformica gracilis shares shortened dorsal tentorial arms, basally angled pedicels, and the pharyngeal gland as apomorphies with other total clade Formicidae. Retained plesiomorphies include mandible shape and features of the prepharynx. Implications of the reconstructed transitions especially for the ant groundplan are critically discussed based on our restricted taxon sampling, emphasizing the crucial information derived from internal anatomy which is applied to deep time for the first time. Based on the falcate mandible in †Gerontoformica and other Aculeata, we present hypotheses for how the shovel-shaped mandibles in crown Formicidae could have evolved. Our results support the notion of †Gerontoformica as ‘generalized’ above-ground predator missing crucial novelties of crown ants which may have helped the latter survive the end-Cretaceous extinction. Our study is an important step for anatomical research on Cretaceous insects and a glimpse into the early evolution of ant heads. Graphical Abstract
化石记录让我们得以一窥当今地球上生物的进化史。我们在克钦琥珀中发现了一个茎类蚂蚁†Gerontoformica gracilis (Barden和Grimaldi, 2014)的标本,其内部头部结构几乎完整地保存了下来,我们使用基于微计算机断层扫描的3D重建记录了这些结构。我们将†Gerontoformica与4个外群分类群和4个现存蚂蚁物种进行比较,利用简约理论和贝叶斯祖先状态重建来识别茎蚁和冠蚁的形态差异和相似性,从而通过头部解剖学的视角提高我们对蚂蚁进化的理解。149个形态特征中有87个是本研究的新发现,几乎都适用于化石。†老年虎蛛与其他虎蛛总分支具有短的背幕臂、基部成角的花梗和咽腺。保留的蛇形包括下颌骨形状和前咽的特征。基于我们的有限分类群采样,重点讨论了重建过渡的意义,特别是对蚂蚁地平面的意义,强调了首次应用于深时间的内部解剖学的关键信息。基于†Gerontoformica和其他Aculeata的镰刀形下颌骨,我们提出了冠Formicidae的铲子形下颌骨进化的假设。我们的研究结果支持†Gerontoformica是一种“广义的”地上捕食者的观点,这种观点缺少了冠蚁的关键新颖性,而这些新颖性可能帮助后者在白垩纪末期的灭绝中幸存下来。我们的研究是白垩纪昆虫解剖学研究的重要一步,也是了解蚂蚁头部早期进化的重要一步。图形抽象
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引用次数: 10
Convergence, Hemiplasy, and Correlated Evolution Impact Morphological Diversity Related to a Web-Less Lifestyle in the Two-Clawed Spiders 趋同、半成型和相关进化影响双爪蜘蛛无网生活方式的形态多样性
IF 3.4 1区 农林科学 Pub Date : 2022-09-01 DOI: 10.1093/isd/ixac020
Guilherme H F Azevedo, T. Bougie, Martín E Carboni, M. Hedin, M. Ramírez
Abstract Traits that independently evolve many times are important for testing hypotheses about correlated evolution and understanding the forces shaping biodiversity. However, population genetics processes can cause hemiplasies (traits determined by genes whose topologies do not match the species tree), leading to a false impression of convergence (homoplasy) and potentially misleading inferences of correlated evolution. Discerning between homoplasies and hemiplasies can be important in cases of rapid radiations and clades with many gene tree incongruences. Here, focusing on two-clawed spiders (Dionycha) and close relatives, we evaluate if the observed distribution of characters related to a web-less lifestyle could be better explained as synapomorphies, homoplasies, or hemiplasies. We find that, although there are several convergences, hemiplasies are also sometimes probable. We discuss how these hemiplasies could affect inferences about correlation and causal relationship of traits. Understanding when and where in the tree of life hemiplasy could have happened is important, preventing false inference of convergent evolution. Furthermore, this understanding can provide alternative hypotheses that can be tested with independent data. Using traits related to the climbing ability of spiders we show that, when hemiplasy is unlikely, adequate model testing can be used to better understand correlated evolution, and propose hypotheses to be tested using controlled behavioral and mechanical experiments. Graphical Abstract
摘要多次独立进化的特征对于检验相关进化的假设和理解塑造生物多样性的力量很重要。然而,群体遗传学过程可能导致异发育(由拓扑结构与物种树不匹配的基因决定的特征),导致趋同的错误印象(同源性),并可能误导相关进化的推断。在快速辐射和具有许多基因树不一致的分支的情况下,区分同源性和半异源性可能很重要。在这里,我们以双爪蜘蛛(Dionycha)和近亲为研究对象,评估观察到的与无网生活方式相关的特征分布是否可以更好地解释为突触形态、同质化或半异变。我们发现,尽管有几种收敛,但有时也可能出现偏侧发育。我们讨论了这些异生现象如何影响关于性状相关性和因果关系的推断。了解生命树中偏瘫发生的时间和地点很重要,可以防止趋同进化的错误推断。此外,这种理解可以提供可以用独立数据检验的替代假设。利用与蜘蛛攀爬能力相关的特征,我们表明,当不太可能出现偏瘫时,可以使用适当的模型测试来更好地理解相关进化,并提出假设,通过受控的行为和机械实验进行测试。图形摘要
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引用次数: 2
How Do Genomic, Mitochondrial, and Morphological Data Contribute to the Linnean Classification of the Porrostomine Net-Winged Beetles (Coleoptera, Lycidae)? 基因组、线粒体和形态学数据如何对波罗托明网翅甲虫(鞘翅目、Lycidae)的林奈分类做出贡献?
IF 3.4 1区 农林科学 Pub Date : 2022-09-01 DOI: 10.1093/isd/ixac023
Dominik Kusy, Michal Motyka, R. Bílková, L. Bocak
The Lycidae genera have seldom been tested with phylogenetic analyses. Therefore, we assembled genomic data to estimate the phylogenetic backbone of the porrostomines, one of Metriorrhynchina’s major clades. Further, mtDNA and morphology were employed to assign 352 analyzed species to genera. We present evidence for the paraphyly of Metriorrhynchus and terminal position of Porrostoma, revise the generic classification, and describe eight genera: Maraiakoreus gen. nov., Kuarhynchus gen. nov., Riedelrhynchus gen. nov., Bundikanus gen. nov., Yamarhynchus gen. nov., Bekorhynchus gen. nov., Sundarhynchus gen. nov., and Isuarhynchus gen. nov. We synonymize Stadenus Waterhouse, 1879, syn. nov., Metriorrhynchoides Kleine, 1923, syn. nov., and Oriomum Bocak, 1999a, syn. nov., to Porrostoma Castelnau, 1838. Next, we propose 75 new combinations and four new species: Bundikanus styskalai sp. nov., Kuarhynchus sisrangensis sp. nov., Maraiakoreus argenteus sp. nov., and Yamarhynchus sinopassensis sp. nov. We identified repeated origins of several external morphological traits earlier used to delimitate genera. Therefore, we prefer concordant evidence from the densely sampled mitochondrial phylogenies and male genitalia. The analyses identify high phylogenetic diversity and species richness in New Guinea, much lower phylogenetic diversity of the Australian continental fauna, and the limited permeability of the Wallacea that resulted in a single porrostomine genus in Asia. We point to the common acceptance of paraphyletic and polyphyletic taxa in the current classification. As a result, taxonomy has not provided expected support for any state-of-the-art evolutionary and zoogeographic studies. The phylogeny, species inventory, and classification of porrostomines set the basis for future evolutionary and zoogeographical studies.
Lycidae属很少进行系统发育分析。因此,我们收集了基因组数据来估计孔虫门的系统发育主干,孔虫门是Metriorhinchina的主要分支之一。此外,利用mtDNA和形态学将352个分析物种划分为属。我们提出了Metriorhynchus和Porrostoma末端位置的旁系性证据,修改了属分类,并描述了八个属:Maraiakorus gen.nov.,Kuarhynchus gen.nv.,Riedelrynchus gen.nov..,Bundikanus gen.no..,Yamarynchus gen.nov。nov.,Metriorhychoides Kleine,1923,syn。nov.和Oriomum Bocak,1999a,syn。11月,致波罗斯托玛·卡斯特尔诺,1838年。接下来,我们提出了75个新组合和4个新种:Bundikanus styskalai sp.nov.、Kuarhynchus sisrangensis sp.nov..、Maraiakoreus Argentieus sp.novs.和Yamarhynchs sinopassinsis sp.nov。我们确定了早期用于划分属的几个外部形态特征的重复起源。因此,我们更喜欢来自密集采样的线粒体系统发育和男性生殖器的一致证据。这些分析表明,新几内亚的系统发育多样性和物种丰富度很高,澳大利亚大陆动物群的系统发育多样化要低得多,而瓦拉科的渗透性有限,导致亚洲只有一个porrostomine属。我们指出,在目前的分类中,副系和多系分类群被普遍接受。因此,分类学并没有为任何最先进的进化和动物地理学研究提供预期的支持。porrostomines的系统发育、物种清单和分类为未来的进化和动物地理学研究奠定了基础。
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引用次数: 1
Whole Genome Sequencing Reveals the Structure of Environment-Associated Divergence in a Broadly Distributed Montane Bumble Bee, Bombus vancouverensis 全基因组测序揭示了广泛分布的山地大黄蜂(Bombus vancouverrensis)与环境相关的分化结构
IF 3.4 1区 农林科学 Pub Date : 2022-09-01 DOI: 10.1093/isd/ixac025
Sam D. Heraghty, Sarthok Rasique Rahman, J. Jackson, Jeffrey D. Lozier
Abstract Broadly distributed species experience divergent abiotic conditions across their ranges that may drive local adaptation. Montane systems where populations are distributed across both latitudinal and elevational gradients are especially likely to produce local adaptation due to spatial variation in multiple abiotic factors, including temperature, oxygen availability, and air density. We use whole-genome resequencing to evaluate the landscape genomics of Bombus vancouverensis Cresson (Hymenoptera: Apidae), a common montane bumble bee that is distributed throughout the western part of North America. Combined statistical approaches revealed several large windows of outlier SNPs with unusual levels of differentiation across the region and indicated that isothermality and elevation were the environmental features most strongly associated with these variants. Genes found within these regions had diverse biological functions, but included neuromuscular function, ion homeostasis, oxidative stress, and hypoxia that could be associated with tolerance of temperature, desiccation, or high elevation conditions.The whole-genome sequencing approach revealed outliers occurred in genome regions with elevated linkage disequilibrium, elevated mean FST, and low intrapopulation nucleotide diversity. Other kinds of structural variations were not widely associated with environmental predictors but did broadly match geographic separation. Results are consistent with other studies suggesting that regions of low recombination may harbor adaptive variation in bumble bees within as well as between species and refine our understanding of candidate genes that could be further investigated as possible targets of selection across the B. vancouverensis range.
广泛分布的物种在其分布范围内经历不同的非生物条件,这些条件可能驱动局部适应。由于温度、氧气有效性和空气密度等多种非生物因素的空间变化,种群分布在纬度和海拔梯度上的山地系统特别可能产生局部适应。本文利用全基因组重测序技术对分布于北美西部的山地大黄蜂——温哥华大黄蜂(Bombus vancouverensis Cresson,膜翅目:蜂科)进行了景观基因组学分析。综合统计方法揭示了几个大窗口的异常snp,在整个地区具有不同寻常的分化水平,并表明等温和海拔是与这些变异最密切相关的环境特征。在这些区域中发现的基因具有多种生物功能,但包括神经肌肉功能、离子稳态、氧化应激和缺氧,这些可能与温度、干燥或高海拔条件的耐受性有关。全基因组测序方法显示,异常值出现在连锁不平衡升高、平均FST升高和种群内核苷酸多样性低的基因组区域。其他类型的结构变化与环境预测因素没有广泛联系,但与地理分离大致相符。结果与其他研究一致,表明低重组区域可能在种内和种间孕育了大黄蜂的适应性变异,并完善了我们对候选基因的理解,这些候选基因可以进一步研究作为温哥华蜜蜂范围内选择的可能目标。
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引用次数: 5
Climatic Niche Conservatism and Ecological Diversification in the Holarctic Cold-Dwelling Butterfly Genus Erebia 北极寒居蝶属Erebia的气候生态位保守性与生态多样性
IF 3.4 1区 农林科学 Pub Date : 2022-08-17 DOI: 10.1093/isd/ixad002
I. Klečková, J. Klečka, Z. Fric, M. Česánek, L. Dutoit, L. Pellissier, Pável Matos‐Maraví
Abstract The diversification of alpine species has been modulated by their climatic niches interacting with changing climatic conditions. The relative roles of climatic niche conservatism promoting geographical speciation and of climatic niche diversification are poorly understood in diverse temperate groups. Here, we investigate the climatic niche evolution in a species rich butterfly genus, Erebia (Dalman, 1816). This Holarctic cold-dwelling genus reaches the highest diversity in European mountains. We generated a nearly complete molecular phylogeny and modeled the climatic niche evolution using geo-referenced occurrence records. We reconstructed the evolution of the climatic niche and tested how the species' climatic niche width changes across the occupied climate gradient and compared two main Erebia clades, the European and the Asian clade. We further explored climatic niche overlaps among species. Our analyses revealed that the evolution of Erebia has been shaped by climatic niche conservatism, supported by a strong phylogenetic signal and niche overlap in sister species, likely promoting allopatric speciation. The European and the Asian clades evolved their climatic niches toward different local optima. In addition, species in the European clade have narrower niches compared to the Asian clade. Contrasts among the clades may be related to regional climate differences, with lower climate seasonality in Europe compared to Central Asia favoring the evolution of narrower niches. Further, adaptive divergence could appear in other traits, such as habitat use, which can be reflected by narrower climatic niches detected in the European clade. Our study extends knowledge about the complexity of evolutionary drivers in temperate insects.
摘要高山物种的多样性受到其气候生态位与气候条件变化相互作用的调节。气候生态位保守性促进地理物种形成和气候生态位多样化的相对作用在不同的温带群体中知之甚少。在这里,我们研究了物种丰富的蝴蝶属Erebia的气候生态位进化(Dalman,1816)。这种全北极寒地居住的属在欧洲山区达到了最高的多样性。我们生成了一个几乎完整的分子系统发育,并使用地理参考发生记录对气候生态位进化进行了建模。我们重建了气候生态位的演变,测试了该物种的气候生态位宽度如何在所占的气候梯度中变化,并比较了两个主要的埃雷比亚分支,即欧洲和亚洲分支。我们进一步探索了物种之间的气候生态位重叠。我们的分析表明,埃雷比亚的进化是由气候生态位保守性决定的,并得到了强大的系统发育信号和姐妹物种生态位重叠的支持,这可能促进了异地物种形成。欧洲和亚洲的分支向不同的局部最优进化其气候生态位。此外,与亚洲分支相比,欧洲分支中的物种具有更窄的生态位。分支之间的对比可能与区域气候差异有关,与中亚相比,欧洲的气候季节性较低,有利于狭窄生态位的进化。此外,适应性差异可能出现在其他特征中,例如栖息地的使用,这可以通过在欧洲分支中检测到的较窄的气候生态位来反映。我们的研究扩展了关于温带昆虫进化驱动因素复杂性的知识。
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引用次数: 4
Utilization of Community Science Data to Explore Habitat Suitability of Basal Termite Genera 利用群落科学资料探讨基底白蚁属生境适宜性
IF 3.4 1区 农林科学 Pub Date : 2022-07-01 DOI: 10.1093/isd/ixac019
Aaron M. Goodman, Jonah J Allen, Jinna Brim, Alessa Codella, Brittney Hahn, Hassan Jojo, Zoila BondocGawa Mafla-Mills, Salka’Tuwa Bondoc Mafla, Agnes Oduro, Megan M. Wilson, J. Ware
Abstract The advent of community-science databases in conjunction with museum specimen locality information has exponentially increased the power and accuracy of ecological niche modeling (ENM). Increased occurrence data has provided colossal potential to understand the distributions of lesser known or endangered species, including arthropods. Although niche modeling of termites has been conducted in the context of invasive and pest species, few studies have been performed to understand the distribution of basal termite genera. Using specimen records from the American Museum of Natural History (AMNH) as well as locality databases, we generated ecological niche models for 12 basal termite species belonging to six genera and three families. We extracted environmental data from the Worldclim 19 bioclimatic dataset v2, along with SoilGrids datasets and generated models using MaxEnt. We chose Optimal models based on partial Receiving Operating characteristic (pROC) and omission rate criterion and determined variable importance using permutation analysis. We also calculated response curves to understand changes in suitability with changes in environmental variables. Optimal models for our 12 termite species ranged in complexity, but no discernible pattern was noted among genera, families, or geographic range. Permutation analysis revealed that habitat suitability is affected predominantly by seasonal or monthly temperature and precipitation variation. Our findings not only highlight the efficacy of largely community-science and museum-based datasets, but our models provide a baseline for predictions of future abundance of lesser-known arthropod species in the face of habitat destruction and climate change. Graphical Abstract
社区科学数据库与博物馆标本位置信息的结合,极大地提高了生态位建模(ENM)的能力和准确性。增加的发生数据为了解鲜为人知或濒危物种(包括节肢动物)的分布提供了巨大的潜力。虽然在入侵和有害物种的背景下进行了白蚁的生态位建模,但很少有研究了解基生白蚁属的分布。利用美国自然历史博物馆(AMNH)的标本记录和当地数据库,建立了隶属于3科6属的12种基础白蚁的生态位模型。我们从Worldclim 19生物气候数据集v2中提取环境数据,以及SoilGrids数据集,并使用MaxEnt生成模型。基于部分接收工作特性(pROC)和遗漏率准则选择最优模型,并利用置换分析确定变量重要性。我们还计算了响应曲线,以了解适应性随环境变量变化的变化。我们的12种白蚁的最优模型的复杂性各不相同,但在属、科或地理范围之间没有明显的模式。生境适宜性主要受季节或月温、降水变化的影响。我们的研究结果不仅突出了社区科学和博物馆数据集的有效性,而且我们的模型为面对栖息地破坏和气候变化的情况下预测未来鲜为人知的节肢动物物种的丰度提供了基线。图形抽象
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引用次数: 1
New Xyleborine (Coleoptera: Curculionidae: Scolytinae) Genus With an Afrotropical-Neotropical Distribution 新木菖蒲属(鞘翅目:菖蒲科),具有热带-新热带分布
IF 3.4 1区 农林科学 Pub Date : 2022-07-01 DOI: 10.1093/isd/ixac016
R. Osborn, Sarah M. Smith, Anthony I Cognato
Abstract Plant-associated arthropods have been shown to cross large oceanic distances on floating plant material and to establish themselves on distant landmasses. Xyleborini (Coleoptera: Curculionidae: Scolytinae) ambrosia beetles occur in forests worldwide and are likely capable of long range dispersal. In less than 20 million years, this group dispersed from Asia to tropical regions of Africa and South America. The phylogeny, taxonomy, and biogeography of one Xyleborus species group which occurs on both continents are reviewed for this study. Based on a well-resolved molecular phylogeny resulting from parsimony, likelihood, and Bayesian analyses of four gene loci, we describe a new monophyletic genus, Xenoxylebora Osborn, Smith & Cognato, gen. nov., for this bicontinental Xyleborus species group with seven Afrotropical and six Neotropical species. Six new species are described: Xenoxylebora pilosa Osborn, Smith & Cognato, sp. nov. from Africa, and Xenoxylebora addenda Osborn, Smith & Cognato, sp. nov., Xenoxylebora calculosa Osborn, Smith & Cognato, sp. nov., Xenoxylebora hystricosa Osborn, Smith & Cognato, sp. nov., Xenoxylebora serrata Osborn, Smith & Cognato, sp. nov., and Xenoxylebora sulcata Osborn, Smith & Cognato, sp. nov., from South America. Seven new combinations from Xyleborus are proposed: Xenoxylebora caudata (Schedl 1957) comb. nov., Xenoxylebora collarti (Eggers 1932) comb. nov., Xenoxylebora perdiligens (Schedl 1937) comb. nov., Xenoxylebora sphenos (Sampson 1912) comb. nov., Xenoxylebora subcrenulata (Eggers 1932) comb. nov., and Xenoxylebora syzygii (Nunberg 1959) comb. nov. from Africa, and Xenoxylebora neosphenos (Schedl 1976) comb. nov. from South America. One new synonym is proposed: Xenoxylebora sphenos (Sampson 1912) = Xyleborus tenellus Schedl 1957 syn. nov. Descriptions, diagnoses, images, and a key to the identification of all 13 species are provided. The sequence of colonization between Africa and South America is uncertain for Xenoxylebora. Prevailing ocean currents and predominant locality patterns observed for other organisms suggest an African Xenoxylebora origin. However, the phylogeny, biogeographical analyses, and a calibrated divergence time suggest a possible South American origin for African Xenoxylebora (2.3 Ma, 95% HDP 4.5–0.6 Ma), which is supported by the occurrence of ocean counter currents between the continents and evidence of dispersal from South America to Africa among some plant and arthropod taxa.
与植物相关的节肢动物已被证明可以依靠漂浮的植物材料跨越遥远的海洋距离,并在遥远的陆地上定居。木蛾(鞘翅目:凤尾蝇科:凤尾蝇科)凤尾虫分布在世界各地的森林中,可能具有远距离传播的能力。在不到2000万年的时间里,这个群体从亚洲分散到非洲和南美洲的热带地区。本文综述了一种分布于两大洲的木耳属植物群的系统发育、分类和生物地理学。基于对4个基因位点的简约性、似然性和贝叶斯分析,我们描述了一个新的单系属,Xenoxylebora Osborn, Smith & Cognato, gen. nov.,这个双大陆木藻物种群有7个非洲热带物种和6个新热带物种。描述了6个新种:来自非洲的piloxlebora Osborn, Smith & Cognato, sp. 11 .和来自非洲的Xenoxylebora Smith & Cognato, Xenoxylebora adenda Osborn, sp. 11 ., Xenoxylebora calculosa Osborn, Smith & Cognato, sp. 11 ., Xenoxylebora hystricosa Osborn, Smith & Cognato, sp. 11 ., Xenoxylebora serrata Osborn, Smith & Cognato, sp. 11 .和来自南美洲的Xenoxylebora sulcata Osborn, Smith & Cognato, sp. 11 .。提出了7个新组合:Xenoxylebora caudata (Schedl 1957) comb。11月,柯氏异叶虫(Eggers, 1932)梳子。11月,perdiligens Xenoxylebora(表1937)梳子。11月,奇异叶虫(Sampson, 1912)梳子。11月,小crenulata Xenoxylebora (Eggers 1932)梳子。nov.,和Xenoxylebora syzygii (Nunberg 1959)梳子。11月产自非洲,和Xenoxylebora neosphenos(1976年计划)梳子。十一月产自南美洲。提出了一个新同义词:Xenoxylebora sphenos (Sampson 1912) = Xyleborus tenellus Schedl 1957 syn11 .提供了所有13种的描述、诊断、图像和识别关键。非洲和南美洲之间的殖民顺序是不确定的Xenoxylebora。在其他生物中观察到的主要洋流和主要地区模式表明非洲蛇尾的起源。然而,系统发育、生物地理分析和校准的分化时间表明,非洲异齿蛇类可能起源于南美洲(2.3 Ma, 95% HDP 4.5-0.6 Ma),大陆间海洋逆流的发生以及一些植物和节肢动物类群从南美洲扩散到非洲的证据支持了这一观点。
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引用次数: 3
Skeletomuscular Atlas and Deep Homology of a Metamorphosing Genitalic Appendage in a Flat-Backed Millipede (Polydesmida: Polydesmidae: Pseudopolydesmus) 平背千足虫一变态生殖器附属物的骨骼肌图谱和深层同源性(多足动物:多足动物科:伪多足动物)
IF 3.4 1区 农林科学 Pub Date : 2022-07-01 DOI: 10.1093/isd/ixac018
Xavier J Zahnle, Megan Ma, J. Bond
Abstract In millipedes, gonopods are male copulatory genitalia derived from walking legs that metamorphose during postembryonic development. The morphology of gonopods is critical for genus and species diagnosis in most taxa. However, the form and function of gonopods vary drastically at the family and ordinal level, making intricate morphological comparison practically impossible. Internal morphology could provide the basis for homologizing morphological elements present in the walking legs and gonopods. Therefore, we used x-ray computed tomography to produce 3D segmentations of Pseudopolydesmus Attems, 1898 (Polydesmida: Polydesmidae) millipedes incorporating two types of morphological elements: skeletal elements and muscles. In addition to imaging the trunk and appendages of an adult male, we imaged the developing gonopod across a series of juvenile male stadia in order to trace the identities of morphological elements. Skeletal elements were homologized, but muscle homologies were limited by the dissimilarity of muscle attachment sites between the walking leg and gonopod. Furthermore, images of juvenile males showed that appendage musculature is totally obliterated once gonopod development begins in the fourth stadium. Due to these limitations, we believe it may be more tractable to homologize gonopod musculature of various millipede taxa to each other than to that of the walking legs. Because distinct genetic developmental networks responsible for walking leg and gonopod patterning have been evolving separately since the common ancestor of gonopod-bearing millipedes (a case of paramorphy), skeletomusculature among gonopods of unrelated millipede taxa is expected to be more similar than that of the gonopod and walking leg within a given millipede lineage.
在千足虫中,性腺是雄性交配生殖器,由胚胎后发育过程中变形的行走腿演变而来。在大多数分类群中,性腺的形态对属和种的诊断至关重要。然而,在科和序的水平上,性腺的形态和功能差异很大,使得复杂的形态比较实际上是不可能的。内部形态可以为行走腿和性腺的形态要素的同源性提供依据。因此,我们使用x射线计算机断层扫描对假足千足虫进行了三维分割,包括两种形态元素:骨骼元素和肌肉。除了对成年雄性的躯干和附属物进行成像外,我们还对一系列幼年雄性的性腺进行了成像,以追踪形态元素的身份。骨骼元素是同源的,但肌肉的同源性受到行走腿和性腺足之间肌肉附着位点的不同的限制。此外,幼年雄性的图像显示,一旦性腺发育开始于第四体育场,附属肌肉组织就完全消失了。由于这些限制,我们认为将不同种类的千足虫的性腺肌肉组织同质化可能比将其与步行腿的肌肉组织同质化更容易处理。由于负责行走腿和性腺模式的不同遗传发育网络自具有性腺足的千足虫的共同祖先(一种副形态的情况)以来一直在分别进化,因此不相关的千足虫类群的性腺动物之间的骨骼肌结构预计比给定千足虫谱系中的性腺动物和行走腿之间的骨骼肌结构更为相似。
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引用次数: 2
Estimating the Drivers of Diversification of Stoneflies Through Time and the Limits of Their Fossil Record 估计石蝇随时间变化的驱动因素及其化石记录的局限性
IF 3.4 1区 农林科学 Pub Date : 2022-07-01 DOI: 10.1093/isd/ixac017
C. Jouault, A. Nel, F. Legendre, F. Condamine
Abstract Deciphering the timing of lineage diversification and extinction has greatly benefited in the last decade from methodological developments in fossil-based analyses. If these advances are increasingly used to study the past dynamics of vertebrates, other taxa such as insects remain relatively neglected. Our understanding of how insect clades waxed and waned or of the impact of major paleoenvironmental changes during their periods of diversification and extinction (mass extinction) are rarely investigated. Here, we compile and analyze the fossil record of Plecoptera (1,742 vetted occurrences) to investigate their genus-level diversification and diversity dynamics using a Bayesian process-based model that incorporates temporal preservation biases. We found that the Permian-Triassic mass extinction has drastically impacted Plecoptera, while the Cretaceous Terrestrial Revolution corresponds with a turnover of plecopteran fauna. We also unveiled three major gaps in the plecopteran fossil record: the Carboniferous-Permian transition, the late Early Cretaceous, and the late Cenomanian to Bartonian, which will need to be further investigated. Based on the life history of extant Plecoptera, we investigate the correlations between their past dynamic and a series of biotic (Red Queen hypothesis) and abiotic (Court Jester hypothesis) factors. These analyses highlight the major role of continental fragmentation in the evolutionary history of stoneflies, which is in line with phylogeny-based biogeographic analyses showing how vicariance drove their diversification. Our study advocates analyzing the fossil record with caution, while attempting to unveil the diversification and extinction periods plus the likely triggers of these past dynamics of diversification.
在过去的十年中,基于化石的分析方法的发展极大地促进了谱系多样化和灭绝时间的破译。如果这些进展越来越多地用于研究脊椎动物过去的动态,其他分类群,如昆虫,相对来说仍然被忽视。我们对昆虫进化枝的兴衰,以及它们在多样化和灭绝(大灭绝)时期主要的古环境变化的影响的理解很少被研究。本文采用贝叶斯过程模型,结合时间保存偏差,对1742种羽翅目的化石记录进行了整理和分析,探讨了羽翅目的属级多样化和多样性动态。我们发现二叠纪-三叠纪大灭绝对翼翅目产生了巨大的影响,而白垩纪陆地革命则与翼翅目动物群的更替相对应。我们还揭示了翅龙化石记录的三个主要空白:石炭纪-二叠纪过渡、早白垩世晚期和晚塞诺曼期到巴顿期,这些空白有待进一步研究。基于现存翼翅目的生活史,研究了其过去的动态与一系列生物(红皇后假说)和非生物(宫廷小丑假说)因素的相关性。这些分析强调了大陆破碎在石蝇进化史中的主要作用,这与基于系统发育的生物地理学分析一致,表明了差异如何推动它们的多样化。我们的研究主张谨慎地分析化石记录,同时试图揭示多样化和灭绝时期以及这些过去多样化动态的可能触发因素。
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引用次数: 7
Mining Ultraconserved Elements From Transcriptome and Genome Data to Explore the Phylogenomics of the Free-living Lice Suborder Psocomorpha (Insecta: Psocodea) 从转录组和基因组数据中挖掘超保守元素以探索自由生活的地衣Psocomorpha亚目(昆虫纲:Psocodea)的系统基因组学
IF 3.4 1区 农林科学 Pub Date : 2022-07-01 DOI: 10.1093/isd/ixac010
Oscar Fernando Saenz Manchola, Ernesto Samacá Sáenz, Stephany Virrueta Herrera, Lorenzo Mario D'Alessio, A. G. García Aldrete, Kevin P. Johnson
Abstract The order Psocodea includes the two historically recognized groups Psocoptera (free-living bark lice) and Phthiraptera (parasitic lice) that were once considered separate orders. Psocodea is divided in three suborders: Trogiomorpha, Troctomorpha, and Psocomorpha, the latter being the largest within the free-living groups. Despite the increasing number of transcriptomes and whole genome sequence (WGS) data available for this group, the relationships among the six known infraorders within Psocomorpha remain unclear. Here, we evaluated the utility of a bait set designed specifically for parasitic lice belonging to suborder Troctomorpha to extract UCE loci from transcriptome and WGS data of 55 bark louse species and explored the phylogenetic relationships within Psocomorpha using these UCE loci markers. Taxon sampling was heavily focused on the families Lachesillidae and Elipsocidae, whose relationships have been problematic in prior phylogenetic studies. We successfully recovered a total of 2,622 UCE loci, with a 40% completeness matrix containing 2,081 UCE loci and an 80% completeness matrix containing 178 UCE loci. The average number of UCE loci recovered for the 55 species was 1,401. The WGS data sets produced a larger number of UCE loci (1,495) on average than the transcriptome data sets (972). Phylogenetic relationships reconstructed with Maximum Likelihood and coalescent-based analysis were concordant regarding the paraphyly of Lachesillidae and Elipsocidae. Branch support values were generally lower in analyses that used a fewer number of loci, even though they had higher matrix completeness. Resumen El orden Psocodea incluye actualmente a dos grupos históricamente reconocidos y que una vez fueron considerados órdenes separados, Psocoptera (piojos de vida libre o de las cortezas) y Phthiraptera (piojos verdaderos). Psocodea está dividido en tres subórdenes: Trogiomorpha, Troctomorpha y Psocomorpha, este último siendo el más grande entre los piojos de vida libre. A pesar de que la cantidad de información disponible sobre transcriptomas y secuenciación del genoma completo (WGS) para este grupo se ha incrementado notablemente en los últimos años, las relaciones filogenéticas dentro de Psocomorpha permanecen poco claras. En este estudio, evaluamos la utilidad de un conjunto de sondas diseñadas específicamente a partir de especies de piojos verdaderos del suborden Troctormorpha, para capturar elementos ultra-conservados (UCE) a partir de las secuencias de transcriptomas y WGS de 55 especies de piojos de las cortezas. Igualmente, exploramos las relaciones filogenéticas dentro de Psocomorpha usando estos marcadores de UCE. El muestreo taxonómico estuvo fuertemente enfocado en las familias Lachesillidae y Elipsocidae, ya que sus relaciones han demostrado ser problemáticas en estudios filogenéticos previos. Como resultado, logramos recuperar exitosamente un total de 2,622 marcadores de UCE, con las matrices de completitud del 40% y 80% conten
摘要PSOCodea秩序包括两个历史上公认的群体PSOCoptera(自由生活的Bark Lice)和PHThiraptera(寄生虫Lice),这两个群体曾经被认为是单独的秩序。PSOCodea分为三个下属:Trogiomorpha、Trocomorpha和PSOComorpha,后者是自由生活群体中最大的。尽管这一群体可获得的转录本和全基因组序列(WGS)数据越来越多,但PSOComorpha内六个已知的次等之间的关系仍然不清楚。在这里,我们评估了一种专门为寄生虫设计的Bait set的用途,该Bait set属于从55种树皮物种的转录本和WGS数据中提取UCE位点的Troctomorpha属,并使用这些UCE位点标记探索了PSOComorpha内部的系统发育关系。分类群取样主要集中在乳蝇科和椭圆体科,它们的关系在以前的系统发育研究中一直存在问题。我们成功地回收了2622个UCE位点,40%的完整性矩阵含有2081个UCE位点,80%的完整性矩阵含有178个UCE位点。为55个物种恢复的平均UCE位点数量为1401个。WGS数据集平均产生的UCE位点(1495个)多于转录组数据集(972个)。用最大似然和基于联合的分析重建的系统发育关系在乳蝇科和椭圆体亚目方面是一致的。在使用少数位点的分析中,分支支持值通常较低,尽管它们的矩阵完整性较高。摘要PSOCodea秩序目前包括两个历史上公认的群体,这两个群体曾经被认为是单独的秩序,即PSOCoptera(自由生活虱子或树皮虱子)和PHTHIRAPTERA(真正的虱子)。PSOCodea分为三个亚目:Trogiomorpha、Trocomorpha和PSOComorpha,后者是自由生活虱子中最大的。尽管近年来,该群体有关转录组和全基因组测序(WGS)的可用信息量显著增加,但PSOComorpha内部的系统发育关系仍不清楚。在这项研究中,我们评估了一组专门从Troctormorpha亚目的真正虱子物种设计的探针在从55种虱子的转录组和WGS序列中捕获超保守元素(UCE)的有用性。同样,我们使用这些UCE标记探索了PSOComorpha内部的系统发育关系。分类取样主要集中在乳蝇科和椭圆体科,因为它们的关系在以前的系统发育研究中被证明是有问题的。结果,我们成功地从UCE中回收了2622个标记,完整性矩阵为40%和80%,分别包含2081个和178个UCE标记。55个物种的平均回收UCE数量为1401个。平均而言,WGS数据集产生的UCE位点(1495个)多于转录组序列(972个)。根据最大似然分析和合并方法重建的系统发育关系与乳蝇科和椭圆体的副翅目是一致的,而在包括较少位点的分析中,即使在完整性矩阵较大的情况下,分支的支持值也通常较低。图形抽象
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引用次数: 2
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Insect Systematics and Diversity
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