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Exploring the stability of sex chromosome systems in Anolis and other iguanas. 探索鬣蜥和其他鬣蜥性染色体系统的稳定性。
IF 2.6 2区 环境科学与生态学 Q2 ECOLOGY Pub Date : 2026-01-30 DOI: 10.1093/evolut/qpag014
Marie Altmanová, Roel M Wouters, Abel Batista, Tomáš Peš, Cristopher A Antúnez-Fonseca, Alejandro J Salguero-Sánchez, Barbora Straková, Lukáš Kubička, Michail Rovatsos, Daniel Ariano-Sánchez, Christian Chinchilla, Michelle Quiroz-Espinoza, Tadeáš Venkrbec, Jasna Vukić, Milan Veselý, Lukáš Kratochvíl

Sex determination was once thought to be unstable in ectothermic vertebrates, but several highly diversified groups of non-avian reptiles, such as iguanas sensu lato (Pleurodonta), appear to have conserved sex chromosomes. However, this statement has been criticized as being based on a parsimonious conclusion from limited sampling. Here, we tested sex chromosome homology in a further 35 species of pleurodont lizards, mainly of the family Anolidae. We demonstrated sex chromosome homology based on the comparison of the gene dosage of X-specific genes by quantitative PCR. Including these new data, the homology of sex chromosomes has so far been supported in 92 out of over 1,200 recently recognized species of Pleurodonta, representing all but one family of this clade. These very conserved sex chromosomes are at least as old as the basal split of Pleurodonta (estimated over 90 MY). In the case of the family Corytophanidae, all genera share different XX/XY sex chromosomes, which are thus over 25 million years old. We discuss the efficiency and limitations of the approach used for tests of the homology of sex chromosomes, as well as the reasons for the evolutionary stability of sex chromosomes in some lineages. We argue that to obtain a complete picture, all extant species should be tested for homology; however, until we reach this ambitious goal, parsimonious estimates in lineages where fragmentary data suggest stability of sex chromosomes, as in Pleurodonta, are substantiated.

性别决定曾经被认为在恒温脊椎动物中是不稳定的,但是一些高度多样化的非鸟类爬行动物群体,如鬣蜥,似乎有保守的性染色体。然而,这一说法被批评为基于有限抽样的简约结论。在这里,我们测试了另外35种胸齿蜥的性染色体同源性,主要是蜥科。通过定量PCR方法比较x特异性基因的基因剂量,证实了性染色体同源性。包括这些新的数据,到目前为止,在1200多个最近发现的侧齿动物物种中,有92个物种的性染色体同源性得到了支持,这些物种代表了除了一个家族之外的所有分支。这些非常保守的性染色体至少与侧齿鱼的基部分裂一样古老(估计超过90年)。以栉水母科为例,所有属都有不同的XX/XY性染色体,因此这些染色体已有2500多万年的历史。我们讨论了用于性染色体同源性测试的方法的效率和局限性,以及性染色体在一些谱系中进化稳定性的原因。我们认为,为了获得一个完整的图像,所有现存的物种都应该进行同源性测试;然而,在我们达到这一雄心勃勃的目标之前,在残片数据表明性染色体稳定性的谱系中,如在Pleurodonta中,吝啬的估计是得到证实的。
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引用次数: 0
Modular reorganization of the trilobite Lonchodomas chaziensis demonstrates mosaic patterns of phenotypic change underlie metamorphosis. 三叶虫chaziensis的模块重组证明了变态背后表型变化的马赛克模式。
IF 2.6 2区 环境科学与生态学 Q2 ECOLOGY Pub Date : 2026-01-30 DOI: 10.1093/evolut/qpag018
Ernesto E Vargas-Parra, Melanie J Hopkins

Some trilobites underwent drastic morphological transformations through their development. The Ordovician trilobite Lonchodomas chaziensis transformed, in a single molt, from a globular protaspid larva to a drastically different adult-like meraspid juvenile. This metamorphosis may be related to a complete shift in lifestyle from a free-floating planktic life-mode into a bottom-dwelling benthic life-mode. By focusing on variation between individuals at distinct ontogenetic stages, shifts in covariation patterns through development can be identified to better understand this transformation. Organisms are composed of parts conceptualized in terms of modules i.e., semi-autonomous packages of highly correlated traits. Modularity is theorized to increase through ontogeny especially in arthropods which undergo metamorphosis. To test this hypothesis, we examine covariation patterns coincident with the restructuring of the trilobite head. Analyses show a decrease in the structure of modularity during metamorphosis and major shifts in the degree of modularity/integration during and preceding transformation. Further, the part which undergoes the most shape change becomes more integrated with other parts. As shape variation is organized in a mosaic fashion through ontogeny, modularity may have played a role in coordinated ontogenetic change among parts leading up to shifts in form and lifestyle occurring during the 'Plankton Revolution' of the early Paleozoic.

有些三叶虫在发育过程中经历了剧烈的形态变化。奥陶系查兹三叶虫Lonchodomas chaziensis在一次蜕皮中,从球状的原蛛幼虫转变为完全不同的成虫样的原蛛幼虫。这种蜕变可能与生活方式的彻底转变有关,从自由漂浮的浮游生物生活模式转变为生活在海底的底栖生物生活模式。通过关注处于不同个体发育阶段的个体之间的变异,可以识别出发育过程中共同变异模式的转变,从而更好地理解这种转变。生物体由以模块为概念的部分组成,即高度相关性状的半自治包。模块化理论通过个体发生增加,特别是在经历变态的节肢动物中。为了验证这一假设,我们研究了与三叶虫头部重组相一致的共变模式。分析表明,在蜕变过程中,模块化结构减少,而在转变过程中和转变之前,模块化/集成度发生了重大变化。此外,经历形状变化最多的部分与其他部分变得更加整合。由于形状变化是通过个体发生以马赛克的方式组织起来的,模块化可能在各部分之间协调的个体发生变化中发挥了作用,导致了早期古生代“浮游生物革命”期间形式和生活方式的转变。
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引用次数: 0
Allometric constraints and the modulation of weapon evolution by mating system in fiddler crabs. 招潮蟹交配系统对异速生长约束和武器进化的调节。
IF 2.6 2区 环境科学与生态学 Q2 ECOLOGY Pub Date : 2026-01-28 DOI: 10.1093/evolut/qpaf217
Cristian L Klunk, Jônatas J Florentino, Daniel S Caetano, Michael S Rosenberg, Alexandre V Palaoro

Sexual selection potentially drives the evolution of exaggerated traits used in intrasexual contests. However, the extent to which mating systems influence weapon morphology remains unclear. In fiddler crab males, an exaggerated claw functions both as a weapon and a signaling tool, varying according to the species' mating system. We examined claw evolution in male fiddler crabs, differentiating between two main mating strategies: (1) males defend their mating burrows (= "burrow"); (2) males do not mate in their own burrows (= "surface"). We measured claw morphological traits and tested whether the mating system affects their evolutionary rates, expecting "burrow" species to exhibit higher evolutionary rates. In general, claw size scales isometrically with body size across species. Both systems showed no correlation between claw elements and mechanical advantage, indicating the necessity of maintaining a conspicuous signaling tool alongside an efficient lever system for grip strength as body size increases. Contrary to predictions, however, "burrow" males exhibited lower evolutionary rates in claw traits than "surface" males, suggesting stronger stabilizing selection. These findings highlight the nuanced effects of sexual selection on male fiddler crab weapon evolution, suggesting that mating systems can modulate evolutionary trajectories, yet functional demands for dual weapon-signal roles constrain claw morphology.

性选择潜在地推动了在无性竞争中使用的夸张特征的进化。然而,交配系统对武器形态的影响程度仍不清楚。在招潮蟹雄性中,一个夸张的爪子既是武器,也是信号工具,根据物种的交配系统而变化。我们研究了雄性招潮蟹的爪子进化,区分了两种主要的交配策略:1。雄性保护它们的交配洞穴(=“洞穴”);2. 雄性不在自己的洞穴(=“表面”)交配。我们测量了爪子的形态特征,并测试了交配制度是否会影响它们的进化速度,预计“穴居”物种会表现出更高的进化速度。一般来说,不同物种的爪子大小与身体大小是等距的。这两个系统都没有显示爪元素和机械优势之间的相关性,这表明随着体型的增加,有必要保持一个明显的信号工具和一个有效的杠杆系统来增强抓握力。然而,与预测相反,“穴居”雄性在爪子特征上的进化速度比“地表”雄性低,这表明稳定选择更强。这些发现强调了性别选择对雄性招潮蟹武器进化的微妙影响,表明交配系统可以调节进化轨迹,但对双重武器信号角色的功能需求限制了爪子形态。
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引用次数: 0
Coevolutionary cycling in allele frequencies and the evolution of virulence. 等位基因频率的共同进化循环与毒力的进化。
IF 2.6 2区 环境科学与生态学 Q2 ECOLOGY Pub Date : 2026-01-28 DOI: 10.1093/evolut/qpaf224
Yoon Soo Kim, Ben Ashby

Coevolutionary cycling in allele frequencies due to negative frequency-dependent selection-sometimes referred to as Red Queen Dynamics-is a key potential outcome of host-parasite coevolution. While many theoretical studies have focused on understanding the consequences of coevolutionary cycling for the evolution of sex and recombination, little is known about the impact of coevolutionary cycling on the evolution of other life history traits. It is therefore currently unknown how coevolutionary cycling in allele frequencies affects the evolution of key disease characteristics, such as virulence. Here, we combine population genetic and quantitative genetic approaches to theoretically determine the impacts of coevolutionary cycling in allele frequencies on the evolution of virulence in a free-living parasite. By varying the level of genetic specificity required for infection while controlling for the average infection rate, we induce coevolutionary cycles and examine their effects on virulence evolution. We show that coevolutionary cycling does indeed have a strong impact on virulence evolution, with more specific infection genetics and higher allelic diversity generally driving larger and more rapid cycles in allele frequencies, leading to selection for higher virulence. Our research provides new fundamental insights into the relationship between coevolutionary cycling and the evolution of virulence.

由于负频率依赖选择导致等位基因频率的共同进化循环——有时被称为红皇后动力学——是宿主-寄生虫共同进化的一个关键潜在结果。虽然许多理论研究都集中在理解共同进化循环对性和重组进化的影响,但对共同进化循环对其他生活史特征进化的影响知之甚少。因此,目前尚不清楚等位基因频率的共同进化循环如何影响关键疾病特征(如毒力)的进化。在这里,我们结合群体遗传学和定量遗传学方法,从理论上确定等位基因频率的共同进化循环对自由生活寄生虫毒力进化的影响。在控制平均感染率的同时,通过改变感染所需的遗传特异性水平,我们诱导了共同进化周期,并研究了它们对毒力进化的影响。我们表明,共同进化循环确实对毒力进化有很强的影响,更具体的感染遗传和更高的等位基因多样性通常会推动等位基因频率更大、更快的循环,从而导致更高毒力的选择。我们的研究为共同进化循环与毒力进化之间的关系提供了新的基本见解。
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引用次数: 0
Reconsidering cytonuclear discordance in the genomic age. 重新考虑基因组时代的细胞核不一致性。
IF 2.6 2区 环境科学与生态学 Q2 ECOLOGY Pub Date : 2026-01-28 DOI: 10.1093/evolut/qpaf201
Drew A Larson, Michael W Itgen, Robert D Denton, Matthew W Hahn

Historically, phylogenetic datasets had relatively few loci but were over-represented for cytoplasmic sequences (mitochondria and chloroplast) because of their ease of amplification and large numbers of informative sites. Under those circumstances, it made sense to contrast individual gene tree topologies obtained from cytoplasmic loci and nuclear loci, with the goal of detecting differences between them-so-called cytonuclear discordance. In the current age of phylogenomics and ubiquitous gene tree discordance among thousands of loci, it is important to distinguish between simply observing discordance between cytoplasmic trees and a species tree inferred from many nuclear loci and identifying the cause of discordance. Here, we examine what inferences one can make from trees representing different genomic compartments. While topological discordance can be caused by multiple factors, the end goal of many studies is to determine whether the compartments have different evolutionary histories: what we refer to as "cytonuclear dissonance." Answering this question is more complex than simply asking whether there is discordance, requiring additional analyses to determine whether genetic exchange has affected only (or mostly) one compartment. Furthermore, even when these histories differ, expectations about why they differ are not always clear. We conclude by pointing to current research and future opportunities that may help to shed light on topological variation across the multiple genomes contained within a single eukaryotic cell.

历史上,系统发育数据集的基因座相对较少,但细胞质序列(线粒体和叶绿体)的基因座过多,因为它们易于扩增和大量的信息位点。在这种情况下,比较从细胞质位点和核位点获得的单个基因树拓扑结构是有意义的,目的是检测它们之间的差异-所谓的细胞核不一致性。在当前的系统基因组学时代,成千上万个基因位点之间普遍存在基因树不一致,区分简单观察细胞质树与从许多核位点推断的物种树之间的不一致和确定不一致的原因是很重要的。在这里,我们研究了人们可以从代表不同基因组区室的树中得出什么推论。虽然拓扑不一致可能由多种因素引起,但许多研究的最终目标是确定这些区室是否具有不同的进化历史:我们称之为“细胞核不和谐”。回答这个问题比简单地问是否存在不一致要复杂得多,需要额外的分析来确定基因交换是否只影响(或主要影响)一个隔间。此外,即使这些历史不同,人们对它们为什么不同的预期也并不总是很清楚。最后,我们指出当前的研究和未来的机会,可能有助于阐明在单个真核细胞中包含的多个基因组的拓扑变异。
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引用次数: 0
Digest: American bats that predominantly feed on plants are more diverse than their generalist relatives. 《文摘》:主要以植物为食的美洲蝙蝠比它们的多面手亲戚更多样化。
IF 2.6 2区 环境科学与生态学 Q2 ECOLOGY Pub Date : 2026-01-28 DOI: 10.1093/evolut/qpaf227
Hernani F M Oliveira, Geraldo B Freire-Jr, Nícholas F Camargo, Daiana C Silva, Sarah S Machado, Joaquim M Silva, Fabrícius M C B Domingos

Omnivory has been hypothesized to be a macroevolutionary sink. A new study by Ochoa-Sanz et al. (2025) tests this hypothesis in Phyllostomidae, a highly ecologically and species-diverse bat family comprising species with different feeding habits, including omnivores and plant specialists. Plant specialists have higher speciation rates than omnivorous bat species, while balanced omnivores have higher speciation rates than plant-predominant ones. Part of the explanation for these differences might be related to the evolution of omnivory during periods of resource scarcity.

杂食被假设为宏观进化的水槽。Ochoa-Sanz等人(2025)的一项新研究在Phyllostomidae中验证了这一假设,Phyllostomidae是一个高度生态和物种多样化的蝙蝠科,由不同食性的物种组成,包括杂食动物和植物专家。植物专门性蝙蝠的物种形成率高于杂食性蝙蝠,而平衡杂食性蝙蝠的物种形成率高于以植物为主的蝙蝠。对这些差异的部分解释可能与资源稀缺时期杂食动物的进化有关。
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引用次数: 0
Can the form of sexual selection explain patterns of static weapon allometry expressed by alternative mating morphotypes? 性选择的形式能否解释由不同的交配形态所表达的静态武器异速发育模式?
IF 2.6 2区 环境科学与生态学 Q2 ECOLOGY Pub Date : 2026-01-28 DOI: 10.1093/evolut/qpaf215
Clint D Kelly

The regime of selection acting on a trait is expected to shape its static allometry. Few studies, however, have quantified the form of sexual selection acting on a trait in the wild to test whether the trait allometrically scales as predicted. Even fewer studies have tested these predictions using males expressing weapon polymorphism as part of their alternative mating strategies. Here, I use field data to test how sexual selection shapes scaling allometries of male weaponry in the Wellington tree wētā (H. crassidens), a male-trimorphic and harem-polygynous insect endemic to New Zealand. Contrary to the prediction that 10th instar males' large weaponry would scale hyperallometrically because it is under direct sexual selection, I found that 10th instar weaponry is not subject to direct sexual selection and scales hypoallometrically. Similarly, neither 8th nor 9th instar male weaponry experiences direct sexual selection, and their weaponry scales hyperallometrically and hypoallometrically, respectively. My study suggests that disentangling competing hypotheses for the evolution of scaling patterns of sexually selected traits must go beyond a simple viability-sexual selection dichotomy by also considering weapon function and the ecological context within which the weapon is used.

作用于性状的选择机制预计会塑造其静态异速生长。然而,很少有研究量化作用于野生性状的性选择形式,以测试该性状是否如预测的那样异速生长。甚至很少有研究将表达武器多态性的雄性作为其替代交配策略的一部分来测试这些预测。在这里,我使用实地数据来测试性选择如何影响威灵顿树wētā (H. crasssidens)雄性武器的缩放异形体,威灵顿树是一种新西兰特有的雄性三型和后肢一夫多妻制昆虫。与预测10龄雄性的大型武器在直接性选择下会呈高异速扩张相反,我发现10龄雄性的大型武器不受直接性选择的影响,而是呈低异速扩张。同样,8龄和9龄雄性武器都没有直接的性选择,它们的武器分别是高异速和低异速。我的研究表明,要解开性选择特征尺度模式进化的相互矛盾的假设,必须超越简单的可行性-性选择二分法,还要考虑武器的功能和使用武器的生态环境。
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引用次数: 0
Overwintering drives rapid adaptation in Drosophila with potential costs to insecticide resistance. 越冬促使果蝇快速适应,但可能会对杀虫剂产生抗性。
IF 2.6 2区 环境科学与生态学 Q2 ECOLOGY Pub Date : 2026-01-28 DOI: 10.1093/evolut/qpaf205
Eric G Prileson, Bianca Campagnari, Catherine I Clare, Amir R Gabidulin, René S Shahmohamadloo, Seth M Rudman

Winter is a formidable challenge for ectotherms that inhabit temperate climates. The extent to which winter conditions drive rapid adaptation, and separately, how selection from novel stressors affects adaptation to winter, remain poorly understood. Here, we use replicate populations of Drosophila melanogaster in a field experiment to test (i) whether winter conditions drive rapid adaptation and (ii) for trade-offs between insecticide resistance and overwintering survival. Following a longitudinal field experiment investigating the evolution of insecticide resistance, we tracked subsequent evolution during an overwintering period. In unexposed control populations, we detected parallel evolutionary shifts indicative of adaptation to winter conditions in multiple traits, including body size and fecundity. Additionally, populations that had evolved insecticide resistance during the growing season were more likely to go extinct than control populations. Further, both control and resistant populations showed patterns of lower resistance following the winter period, suggestive of a trade-off between overwintering success and insecticide resistance. Rapid evolutionary responses to winter conditions, and potential costs of resistance, provide important context for understanding overwintering performance in temperate insects with implications for pest management and ecosystem services.

对于生活在温带气候的变温动物来说,冬天是一个巨大的挑战。冬季条件在多大程度上推动了快速适应,以及从新的压力源中选择如何影响对冬季的适应,人们仍然知之甚少。在这里,我们使用重复种群的黑腹果蝇进行田间实验,以测试i)冬季条件是否推动快速适应ii)杀虫剂抗性与越冬生存之间的权衡。在调查杀虫剂抗性演变的纵向田间试验之后,我们跟踪了越冬期间的后续演变。在未暴露的对照种群中,我们发现了平行的进化变化,表明多种特征(包括体型和繁殖力)适应冬季条件。此外,在生长季节进化出杀虫剂抗性的种群比对照种群更容易灭绝。此外,对照种群和抗性种群在冬季后均表现出较低的抗性模式,这表明越冬成功与杀虫剂抗性之间存在权衡关系。对冬季条件的快速进化反应和抗性的潜在代价,为了解温带昆虫的越冬行为提供了重要背景,对有害生物管理和生态系统服务具有重要意义。
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引用次数: 0
Tetrapod vocal evolution reveals faster rates and higher-pitched sounds for mammals. 四足动物的声音进化揭示了哺乳动物更快的速度和更高的音调。
IF 2.6 2区 环境科学与生态学 Q2 ECOLOGY Pub Date : 2026-01-28 DOI: 10.1093/evolut/qpaf209
Matías I Muñoz, Myriam Marsot, Jacintha Ellers, Wouter Halfwerk

Using the voice to produce sound is a widespread form of communication and plays an important role across diverse species and contexts. Variation in the rate and mode of sound production has been extensively studied within orders or classes, but understanding vocal signal evolution ultimately requires comparison across all major lineages involved. Here, we used phylogenetic comparative methods to investigate the evolution of dominant frequency and its association with body mass across a set of 873 species of mammals, birds, and frogs. Our results show that all vocal systems share the same general feature of the negative allometric relationship between body mass and dominant frequency, but that mammals clearly deviate compared to frogs and birds. We found mammals to vocalize at much higher frequencies and their signals evolved four- to sixfold faster compared to other tetrapod clades. Although all three groups strongly rely on vocal communication, our findings show that only mammals have extensively explored the spectral acoustic space. We argue that such high vocal diversity of mammals is made possible by their unique hearing system, and discuss the functional drivers that allowed their shared ancestors to evolve a richer array of frequencies than other tetrapods.

用声音发声是一种广泛的交流方式,在不同的物种和环境中发挥着重要作用。声音产生的速率和模式的变化已经在目或类中得到了广泛的研究,但理解声音信号的进化最终需要对所有主要谱系进行比较。本文采用系统发育比较方法,研究了873种哺乳动物、鸟类和青蛙的显性频率的进化及其与体重的关系。我们的研究结果表明,所有的发声系统都具有相同的总体特征,即体重和主导频率之间的负异速关系,但与青蛙和鸟类相比,哺乳动物明显偏离了这一特征。我们发现哺乳动物的发声频率要高得多,它们的信号进化速度比其他四足动物快4到6倍。尽管这三种动物都强烈依赖于声音交流,但我们的研究结果表明,只有哺乳动物才广泛地探索了光谱声学空间。我们认为,哺乳动物如此高的声音多样性是由它们独特的听觉系统造成的,并讨论了使它们共同的祖先进化出比其他四足动物更丰富的频率阵列的功能驱动因素。
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引用次数: 0
Repeated, irreversible evolution of ant-following behavior across Neotropical avian families. 新热带鸟类家族中蚂蚁跟随行为的重复、不可逆进化。
IF 2.6 2区 环境科学与生态学 Q2 ECOLOGY Pub Date : 2026-01-28 DOI: 10.1093/evolut/qpaf214
William Sweet, Justin Baldwin, Juan Pablo Gomez, Henry Pollock, Ari E Martínez

Ecological specialization is a result of the interplay between ecological and evolutionary processes. One iconic ecological specialization of the Neotropics involves birds that follow army ant swarms in feeding groups. Prior work has focused on a single avian family, the Neotropical antbirds (Thamnophilidae), but over a century of fieldwork has now revealed that ant-following occurs in hundreds of distantly related birds. To understand the relative contributions of shared ancestry and ecological specialization in the evolution of ant-following, we compiled a database of all Neotropical ant-following birds (n = 472 species) and their degree of specialization on army ants, and tested if (1) ant-following becomes increasingly specialized through evolutionary time and (2) ecomorphological functional traits predict ant-following behavior. Ancestral state reconstruction revealed that specialized ant-following evolved independently in 8 clades and 4 families of Neotropical birds (Antbirds: Thamnophilidae, Ovenbirds: Furnariidae, Tanagers: Thraupidae, and Cuckoos: Cuculidae). Ant-following behavior was highly conserved phylogenetically (Pagel's λ = 0.97), and specialized clades evolved from less specialized ancestors, with few evolutionary reversals. In contrast, ecomorphological traits poorly predicted the level of ant-following specialization across species. Our results suggest increasing specialization on army ants is governed by niche conservatism, not ecological specialization.

生态专门化是生态过程和进化过程相互作用的结果。新热带地区一个标志性的生态专一性涉及到跟随军蚁群觅食的鸟类。先前的研究主要集中在一个单一的鸟类家族,新热带蚁鸟(Thamnophilidae),但是一个多世纪的田野调查现在已经揭示了蚂蚁跟随现象发生在数百种远亲鸟类中。为了了解共同祖先和生态专一化在蚁步进化中的相对贡献,我们编制了所有新热带蚁步鸟类(n = 472种)的数据库及其对军蚁的专一化程度,并测试了:1)蚁步随着进化时间的推移变得越来越专一,2)生态形态学功能特征是否预测了蚁步行为。祖先状态重建表明,新热带鸟类的8个分支和4个科(蚁鸟:蚁巢科,炉鸟:蚁巢科,雀鸟:鸫科,杜鹃科)独立进化出了专门的蚁步。蚂蚁跟随行为在系统发育上是高度保守的(Pagel λ = 0.97),特化的进化枝是从不那么特化的祖先进化而来的,很少有进化逆转。相比之下,生态形态特征很难预测跨物种的蚂蚁跟随专业化水平。我们的研究结果表明,军蚁的专业化程度是由生态位保守主义控制的,而不是生态专业化。
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引用次数: 0
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Evolution
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