Biology subjects

Hoehna, S.

Publications and source records attributed to Hoehna, S..

At least 19 recordsLinked to original sources

bioRxiv2026

How Robust are Multispecies Coalescent Species Delimitations in Taxonomically Complex Systems? A Genomic Assessment Using Mediterranean Tethya Sponges

Reliable species delimitation underpins biodiversity assessment but remains difficult for organisms with plastic morphology and few diagnostic characters. Multispecies coalescent (MSC) methods can delimit species from genomic data, yet they are rarely tested in taxonomically complex, marine invertebrate groups where they are arguably most needed. We used the three Mediterranean species of the genus Tethya, a rare, well-characterised system within the otherwise taxonomically difficult phylum Porifera--distinguished by multiple independent morphological and ecological characters--to evaluate how robust MSC-based delimitation is in such groups. Analysing 64 single-copy nuclear loci in BEAST2 and BPP, we compared constrained, hypothesis-testing approaches (BFD*, BFdriver, A10) with freer, heuristic ones (SPEEDEMON, A11), and examined their sensitivity to data type, clock model, priors, and the species-collapse threshold. All methods recovered the three recognised Mediterranean species, but the resolution of within-lineage structure was method-dependent. The hypothesis-testing approaches consistently supported six lineages, robustly across data types and model assumptions, whereas the heuristic approaches proved less stable. Configurations without a priori species hypotheses often failed to converge or were computationally intractable, a problem compounded by the relaxed clock. In SPEEDEMON the outcome changed with the collapse threshold. Because our system lacks an independent reference point to calibrate this threshold, any delimitation based on it is poorly constrained. We conclude that constrained, hypothesis-testing delimitation is the most robust and reproducible MSC approach, yielding a quantitative, model-based hypothesis that can be weighed against other lines of evidence to inform taxonomic decisions. By clarifying how these methods behave and how their outcomes should be interpreted, our study offers a practical guide for researchers working on comparably complex systems.

evolutionary biology↗

bioRxiv2025

Impact of Partition Models on Phylogenetic Inference and Divergence Times of Lampyridae from Mitochondrial Genomes

Mitochondrial genomes are frequently used for phylogenetic inference due to their availability and cost-efficient sequencing. In most mitogenomic phylogenetic analyses, only the two ribosomal RNA and 13 protein coding genes are used. For such multi-locus datasets it has long been established that appropriate data partition models, e.g., partitioning by gene type and/or codon position, are necessary for robust phylogenetic inference. While most studies focused on the impact of partition models on the tree topology, little is known about the impact on divergence time estimation. Furthermore, although modeling among site rate variation within a partition is common practice, the extent of substitution rate variation among partitions is less explored. Here we study the impact of four partition model strategies: (i) no subdivision of the data (uniform), (ii) partitioning by gene, (iii) partitioning by codon position, and (iv) partitioning by both gene and codon position (combined). We explore the impact of partition models on divergence time estimation in fireflies (Coleoptera: Lampyridae). To this end, we sequenced mitochondrial genomes from 22 firefly species from Europe and Central America and combined these with 82 published mitochondrial genomes. Our results represent the most comprehensive time-calibrated phylogeny of fireflies to day. While the partition models had an impact on the inferred tree topology, the divergence times were almost not affected. Furthermore, we observed up to 3-fold substitution rate variation across genes and additionally more than to 10-fold substitution rate variation across codon positions. Our study gives insights into best practices of performing partitioned-data time-calibrated phylogenetic inference from mitochondrial genomes and multi-locus datasets in general.

evolutionary biology↗

bioRxiv2025

A covarion model for phylogenetic estimation using discrete morphological datasets

AbstractThe rate of evolution of a single morphological character is not homogeneous across the phylogeny and this rate heterogeneity varies between morphological characters. However, traditional models of morphological character evolution often assume that all characters evolve according to a time-homogeneous Markov process, which applies uniformly across the entire phylogeny. While models incorporating amongcharacter rate variation alleviate the assumption of the same rate for all characters, they still fail to address lineage-specific rate variation for individual characters. The covarion model, originally developed for molecular data to model the invariability of some sites for parts of the phylogeny, provides a promising framework for addressing this issue in morphological phylogenetics. In this study, we extend the covarion model in RevBayes to morphological character evolution, which we call the covariomorph model, and apply it to a diverse range of morphological datasets. Our covariomorph model utilizes multiple rate categories derived from a discretized probability distribution, which scales rate matrices accordingly. Characters are allowed to evolve within any of these rate categories, with the possibility of switching between rate categories during the evolutionary process. We verified our implementation of the covariomorph model with the help of simulations. Additionally, we examined 164 empirical datasets, finding patterns of rate heterogeneity compatible with covarion-like dynamics in approximately half of them. Upon further examination of two focal datasets that exhibited covarion-like rate variation, we found that the covariomorph model provides a more nuanced approach to incorporate rate variation across lineages, significantly affecting the resulting tree topology and branch lengths compared to traditional models. The observed sensitivity of branch lengths to model choice underscores potential implications of this approach for divergence time estimation and evolutionary rate calculations. By accounting for lineageand character-specific rate shifts, the covariomorph model offers a robust framework to improve the accuracy of morphological phylogenetic inference.

evolutionary biology↗

bioRxiv2025

Filling the gap in the Neotropics: new firefly species provide insights into Central American biodiversity

Tropical ecosystems, particularly in the Neotropics, remain vastly understudied, especially regarding the biodiversity of many taxa such as fireflies. This study investigates the biodiversity and phylogenetic relationships of fireflies in Guatemala, focusing on the genera Photinus, Photuris, and Bicellonycha. Using genetic barcoding we generate time-calibrated phylogenetic trees, revealing deep divergent times between Guatemalan and North American fireflies and potentially multiple colonialization events. Using morphometric information, taxonomic keys and species delimitations models (mPTP) we identified four new Photinus species: P. hunahpu, P. helodermensis, P. schusteri, and P. semetabajense. This work highlights the significant firefly biodiversity in Nuclear Central America and the potential for future species discovery. Biodiversity research is necessary for conservation efforts of key ecosystems and will broaden our understanding of general evolutionary processes.

evolutionary biology↗

bioRxiv2024

On the Mkv Model with Among-Character Rate Variation

Models used in likelihood-based morphological phylogenetics often adapt molecular phylogenetics models to the specificities of morphological data. Such is the case for the widely used Mkv model-- which introduces an acquisition bias correction for sampling only characters that are observed to be variable--and for models of among-character rate variation (ACRV), routinely applied by researchers to relax the equal-rates assumption of Mkv. However, the interaction between variable character acquisition bias and ACRV has never been explored before. We demonstrate that there are two distinct approaches to condition the likelihood on variable characters when there is ACRV, and we call them joint and marginal acquisition bias. Far from being just a trivial mathematical detail, we show that the way in which the variable character conditional likelihood is calculated results in different assumptions about how rate variation is distributed in morphological datasets. Simulations demonstrate that tree length and amount of ACRV in the data are systematically biased when conditioning on variable characters differently from how the data was simulated. Moreover, an empirical case study with extant and extinct taxa reveals a potential impact not only on the estimation of branch lengths, but also of phylogenetic relationships. We recommend the use of the marginal acquisition bias approach for morphological datasets modeled with ACRV. Finally, we urge developers of phylogenetic software to clarify which acquisition bias correction is implemented for both estimation and simulation, and we discuss the implications of our findings on modeling variable characters for the future of morphological phylogenetics.

evolutionary biology↗

bioRxiv2024

Mixture Models for Dating with Confidence

Robust estimation of divergence times is commonly performed using Bayesian inference with relaxed clock models. The specific choice of relaxed clock model and tree prior model can impact divergence time estimates, thus necessitating model selection among alternative models. The common approach is to select a model based on Bayes factors estimated via computational demanding approaches such as stepping stone sampling. Here we explore an alternative approach: mixture models that analytically integrate over all candidate models. Our mixture model approach only requires one Markov chain Monte Carlo analysis to both estimate the parameters of interest (e.g., the time-calibrated phylogeny) and to compute model posterior probabilities. We demonstrate the application of our mixture model approach using three relaxed clock models (uncorrelated exponential, uncorrelated lognormal and independent gamma rates) combined with three tree prior models (constant-rates pure birth process, constant-rate birth-death process and piecewise-constant birth-death process) and mitochondrial genome dataset of Crocodylia. We calibrate the phylogeny using well-defined fossil node calibrations. Our results show that Bayes factors estimated using stepping stone sampling are unreliable due to noise in repeated analyses while our analytical mixture model approach shows higher precision and robustness. Thus, divergence time estimates under our mixture model are comparably robust as previous relaxed clock approaches but model selection is significantly faster and avoids marginal likelihood estimation. Finally, our time-calibrated phylogeny of Crocodylia presents a robust benchmark for further studies in the group.

evolutionary biology↗

bioRxiv2024

A 20+ Ma old enamel proteome from Canada's High Arctic reveals diversification of Rhinocerotidae in the middle Eocene-Oligocene

In the past decade, ancient protein sequences have emerged as a valuable source of data for deep-time phylogenetic inference. Still, the recovery of protein sequences providing novel phylogenetic insights does not exceed 3.7 Ma (Pliocene). Here, we push this boundary back to 21-24 Ma (early Miocene), by retrieving enamel protein sequences of an early-diverging rhinocerotid (Epiaceratherium sp. - CMNF-59632) from the Canadian High Arctic. We recover partial sequences of seven enamel proteins (AHSG, ALB, AMBN, AMELX, AMTN, ENAM, MMP20) and over 1000 peptide-spectrum matches, spanning over at least 251 amino acids. Authentic endogeneity of these sequences is supported by indicators of protein damage, including several spontaneous and irreversible post-translational modifications accumulated during prolonged diagenesis and reaching near-complete occupancy at many sites. Bayesian tip-dating, across 15 extant and extinct perissodactyl taxa, places the divergence time of CMNF-59632 in the middle Eocene-Oligocene, and identifies a later divergence time for Elasmotheriinae in the Oligocene. The finding weakens alternative models suggesting a deep basal split between Elasmotheriinae and Rhinocerotinae. This divergence time of CMNF-59632 coincides with a phase of high diversification of rhinocerotids, and supports a Eurasian origin of this clade in the late Eocene or Oligocene. The findings are consistent with previous hypotheses on the origin of the enigmatic fauna of the Haughton crater, which, in spite of their considerable degree of endemism, also display similarity to distant Eurasian faunas. Our findings demonstrate the potential of palaeoproteomics in obtaining phylogenetic information from a specimen that is ten times older than any sample from which endogenous DNA has been obtained.

paleontology↗

bioRxiv2024

Sex-biased migration and demographic history of the big European firefly Lampyris noctiluca

Differential dispersion between the sexes can impact population structure and connectivity between populations, which in turn, can have an effect on the evolvability capability of a species. Here we explored the demographic history of the big European firefly, Lampyris noctiluca, which exhibits female neoteny. L. noctiluca distribution extends throughout Europe, but nothing is known on how its colonization processes. To investigate this, we produced the first Lampyris genome (653Mb), including an IsoSeq annotation and the identification of the X chromosome. We collected 115 individuals from six populations of L. noctiluca (Finland to Italy) and generated whole genome re-sequencing for each individual. We inferred several population expansions and bottlenecks throughout the Pleistocene that correlate with glaciation events. Surprisingly, we uncovered strong population structure and low gene-flow. We reject a stepwise, south to north, colonization history scenario and instead uncovered a complex demographic history with a putative eastern European origin. Analyzing the evolutionary history of the mitochondrial genome as well as X-linked and autosomal loci, we found evidence of a maternal colonialization of Germany, putatively from a western European population, followed by male-only migration from south of the Alps (Italy). Overall, investigating the demographic history and colonization patterns of a species should form part of an integrative approach of biodiversity research. Our results provide evidence of sex-biased migration which is important to consider for demographic, biogeographic and species delimitation studies.

evolutionary biology↗

bioRxiv2024

Assessing the Adequacy of Morphological Models used in Palaeobiology

Reconstructing the evolutionary history of different groups of organisms provides insight into how life originated and diversified on Earth. Phylogenetic trees are commonly used to estimate this evolutionary history, providing a hypothesis of the events. Within Bayesian phylogenetics a major step in estimating a tree is in choosing an appropriate model of character evolution. In the case of most extinct species, our only source of information to decipher their phylogenetic relationships is through the morphology of fossils. We therefore use a model of morphological character evolution, the most common of which being the Mk Lewis model. While it is frequently used in palaeobiology, it is not known whether the simple Mk substitution model, or any extensions to it, provide a sufficiently good description of the process of morphological evolution. To determine whether or not the Mk model is appropriate for fossil data we used posterior predictive simulations, a model adequacy approach, to estimate absolute fit of the model to morphological data sets. We first investigate the impact that different versions of the Mk model have on key parameter estimates using tetrapod data sets. We show that choice of substitution model has an impact on both topology and branch lengths, highlighting the importance of model choice. Next, we use simulations to investigate the power of posterior predictive simulations for morphology. Having validated this approach we show that current variations of the Mk model are in fact performing adequately in capturing the evolutionary dynamics that generated our data. We do not find any preference for a particular model extension across multiple data sets, indicating that there is no one size fits all when it comes to morphological data and that careful consideration should be given to choosing models of discrete character evolution. By using suitable models of character evolution, we can increase our confidence in our phylogenetic estimates, which should in turn allow us to gain more accurate insights into the evolutionary history of both extinct and extant taxa.

paleontology↗

bioRxiv2022

ACDC: Analysis of Congruent Diversification Classes

1. Diversification rates inferred from phylogenies are not identifiable. There are infinitely many combinations of speciation and extinction rate functions that have the exact same likelihood score for a given phylogeny, building a congruence class. The specific shape and characteristics of such congruence classes have not yet been studied. Whether speciation and extinction rate functions within a congruence class share common features is also not known. 2. Instead of striving to make the diversification rates identifiable, we can embrace their inherent non-identifiable nature. We use two different approaches to explore a congruence class: (i) testing of specific alternative hypotheses, and (ii) randomly sampling alternative rate function within the congruence class. 3. Our methods are implemented in the open-source R package ACDC (https://github.com/afmagee/ACDC). ACDC provides a flexible approach to explore the congruence class and provides summaries of rate functions within a congruence class. The summaries can highlight common trends, i.e. increasing, flat or decreasing rates. Although there are infinitely many equally likely diversification rate functions, these can share common features. ACDC can be used to assess if diversification rate patterns are robust despite non-identifiability. In our example, we clearly identify three phases of diversification rate changes that are common among all models in the congruence class. Thus, congruence classes are not necessarily a problem for studying historical patterns of biodiversity from phylogenies.

evolutionary biology↗

bioRxiv2021

Polymorphism-aware estimation of species trees and evolutionary forces from genomic sequences with RevBayes

O_LIThe availability of population genomic data through new sequencing technologies gives unprecedented opportunities for estimating important evolutionary forces such as genetic drift, selection, and mutation biases across organisms. Yet, analytical methods that can handle polymorphisms jointly with sequence divergence across species are rare and not easily accessible to empiricists. C_LIO_LIWe implemented polymorphism-aware phylogenetic models (PoMos), an alternative approach for species tree estimation, in the Bayesian phylogenetic software RevBayes. PoMos naturally account for incomplete lineage sorting, which is known to cause difficulties for phylogenetic inference in species radiations, and scale well with genome-wide data. Simultaneously, PoMos can estimate mutation and selection biases. C_LIO_LIWe have applied our methods to resolve the complex phylogenetic relationships of a young radiation of Chorthippus grasshoppers, based on coding sequences. In addition to establishing a well-supported species tree, we found a mutation bias favoring AT alleles and selection bias promoting the fixation of GC alleles, the latter consistent with GC-biased gene conversion. The selection bias is two orders of magnitude lower than genetic drift, validating the critical role of nearly neutral evolutionary processes in species radiation. C_LIO_LIPoMos offer a wide range of models to reconstruct phylogenies and can be easily combined with existing models in RevBayes -- e.g., relaxed clock and divergence time estimation -- offering new insights into the evolutionary processes underlying molecular evolution and, ultimately, species diversification. C_LI

bioinformatics↗

bioRxiv2021

A Time-calibrated Firefly (Coleoptera: Lampyridae) Phylogeny: Using Genomic Data for Divergence Time Estimation

Fireflies (Coleoptera: Lampyridae) consist of over 2,000 described extant species. A well-resolved phylogeny of fireflies is important for the study of their population genetics, bioluminescence, evolution, and conservation. We used a recently published anchored hybrid enrichment dataset (AHE; 436 loci for 88 Lampyridae species and 10 outgroup species) and state-of-the-art statistical methods (the fossilized birth-death-range process implemented in a Bayesian framework) to estimate a time-calibrated phylogeny of Lampyridae. Unfortunately, estimating calibrated phylogenies using AHE and the latest and most robust time-calibration strategies is not possible because of computational constraints. As a solution, we subset the full dataset by applying three different strategies: (i) using the most complete loci, (ii) using the most homogeneous loci, and (iii) using the loci with the highest accuracy to infer the well established Photinus clade. The estimated topology using the three data subsets agreed on almost all major clades and only showed minor discordance within less supported nodes. The estimated divergence times overlapped for all nodes that are shared between the topologies. Thus, divergence time estimation is robust as long as the topology inference is robust and any well selected data subset suffices. Additionally, we observed an un-expected amount of gene tree discordance between the 436 AHE loci. Our assessment of model adequacy showed that standard phylogenetic substitution models are not adequate for any of the 436 AHE loci which is likely to bias phylogenetic inferences. We performed a simulation study to explore the impact of (a) incomplete lineage sorting, (b) uniformly distributed and systematic missing data, and (c) systematic bias in the position of highly variable and conserved sites. For our simulated data, we observed less gene tree variation which shows that the empirically observed amount of gene tree discordance for the AHE dataset is unexpected and needs further investigation.

evolutionary biology↗

bioRxiv2021

Improved resolution of recalcitrant nodes in the animal phylogeny through the analysis of genome gene content and morphology

An accurate phylogeny of animals is needed to clarify their evolution, ecology, and impact on shaping the biosphere. Although datasets of several hundred thousand amino acids are nowadays routinely used to test phylogenetic hypotheses, key deep nodes in the metazoan tree remain unresolved: the root of animals, the root of Bilateria, and the monophyly of Deuterostomia. Instead of using the standard approach of amino acid datasets, we performed analyses of newly assembled genome gene content and morphological datasets to investigate these recalcitrant nodes in the phylogeny of animals. We explored extensively the choices for assembling the genome gene content dataset and model choices of morphological analyses. Our results are robust to these choices and provide additional insights into the early evolution of animals, they are consistent with sponges as the sister group of all the other animals, the worm-like bilaterian lineage Xenacoelomorpha as the sister group of the other Bilateria, and tentatively support monophyletic Deuterostomia.

evolutionary biology↗

bioRxiv2021

The rise of grasslands is linked to atmospheric CO2 decline in the late Palaeogene

Grasslands are predicted to experience a major biodiversity change by the year 2100 in part due to recent and projected increases in atmospheric CO2 concentration. A better understanding of how grasslands have responded to past environmental changes will help predict the outcome of current and future environmental changes. Here, we explore the relationship between past atmospheric CO2 and temperature fluctuations and the shifts in diversification rate of grasses (Poaceae) and daisies (Asteraceae), two exceptionally species-rich grassland families ([~]11,000 and [~]23,000 species, respectively). To this end, we developed a novel Bayesian approach that simultaneously estimates diversification-rates through time from time-calibrated phylogenies and correlations between environmental variables and diversification rates. Additionally, we developed a new statistical approach that incorporates the information of the distribution of missing species in the phylogeny. We found strong evidence supporting a simultaneous increase in diversification rates for daisies and grasses after the most significant reduction of atmospheric CO2 in the Cenozoic ([~]34 Mya). The fluctuations of paleo-temperatures, however, appear not to have had a significant relationship with the diversification of these grassland families. Overall, our results shed new light on our understanding of the origin of grasslands in the context of past environmental changes.

evolutionary biology↗

bioRxiv2021

Convergence Assessment for Bayesian Phylogenetic Analysis using MCMC simulation

O_LIPosterior distributions are commonly approximated by samples produced from a Markov chain Monte Carlo (MCMC) simulation. Every MCMC simulation has to be checked for convergence, i.e., that sufficiently many samples have been obtained and that these samples indeed represent the true posterior distribution. C_LIO_LIHere we develop and test different approaches for convergence assessment in phylogenetics. We analytically derive a threshold for a minimum effective sample size (ESS) of 625. We observe that only the initial sequence estimator provides robust ESS estimates for common types of MCMC simulations (autocorrelated samples, adaptive MCMC, Metropolis-Coupled MCMC). We show that standard ESS computation can be applied to phylogenetic trees if the tree samples are converted into traces of absence/presence of splits. C_LIO_LIConvergence in distribution between replicated MCMC runs can be assessed with the Kolmogorov-Smirnov test. The commonly used potential scale reduction factor (PSRF) is biased when applied to skewed posterior distribution. Additionally, we analytically derive the expected difference between split frequencies (EDSF) and show that it depends on the true frequency of a split. Hence, the average standard deviation of split frequencies is too simplistic and the EDSF should be used instead to check for convergence in split frequencies. C_LIO_LIWe implemented the methods described here in the open-source R package Convenience (https://github.com/lfabreti/convenience), which allows users to easily test for convergence using output from standard phylogenetic inference software. C_LI

bioinformatics↗

bioRxiv2020

Stepwise Bayesian Phylogenetic Inference

The ideal approach to Bayesian phylogenetic inference is to estimate all parameters of interest jointly in a single hierarchical model. However, this is often not feasible in practice due to the high computational cost that would be incurred. Instead, phylogenetic pipelines generally consist of chained analyses, whereby a single point estimate from a given analysis is used as input for the next analysis in the chain (e.g., a single multiple sequence alignment is used to estimate a gene tree). In this framework, uncertainty is not propagated from step to step in the chain, which can lead to inaccurate or spuriously certain results. Here, we formally develop and test the stepwise approach to Bayesian inference, which uses importance sampling to generate observations for the next step of an analysis pipeline from the posterior produced in the previous step. We show that this approach is identical to the joint approach given sufficient information in the data and in the importance sample. This is demonstrated using both a toy example and an analysis pipeline for inferring divergence times using a relaxed clock model. The stepwise approach presented here not only accounts for uncertainty between analysis steps, but also allows for greater flexibility in program choice (and hence model availability) and can be more computationally efficient than the traditional joint approach when multiple models are being tested.

evolutionary biology↗

bioRxiv2020

Relative time constraints improve molecular dating

Dating the tree of life is central to understanding the evolution of life on Earth. Molecular clocks calibrated with fossils represent the state of the art for inferring the ages of major groups. Yet, other information on the timing of species diversification can be used to date the tree of life. For example, horizontal gene transfer events and ancient coevolutionary relationships such as (endo)symbioses can imply temporal relationships between two nodes in a phylogeny (Davin et al. 2018). These alternative sources of temporal constraints can be particularly helpful when the geological record is sparse, e.g. for microorganisms, which represent the vast majority of extant and extinct biodiversity. Here, we present a new method to combine fossil calibrations and relative age constraints to estimate chronograms. We provide an implementation of relative age constraints in RevBayes (Hohna et al. 2016) that can be combined in a modular manner with the wide range of molecular dating methods available in the software. We use both realistic simulations and empirical datasets of 40 Cyanobacteria and 62 Archaea to evaluate our method. We show that the combination of relative age constraints with fossil calibrations significantly improves the estimation of node ages.

evolutionary biology↗

bioRxiv2019

On the inference of a southern origin of the North American firefly Photinus pyralis

The firefly Photinus pyralis inhabits a wide range of latitudinal and ecological niches, with populations living from temperate to tropical habitats. Its ample geographic distribution makes this species an ideal system for the study of local adaptation and demographic inference of wild populations. Therefore, in this study we modelled and inferred different demographic scenarios for North American populations of P. pyralis, collected from Texas to New Jersey. To do this, we used a combination of ABC techniques (for multi-population/colonization analyses), and likelihood inference (dadi) for single-population demographic inference, which proved useful with our RAD data.We uncovered that the most ancestral North American population lays in Texas, which further colonized the Central region of the US and more recently the North Eastern coast. Our study confidently rejects a demographic scenario where the North Eastern populations colonized more southern populations until reaching Texas. Our results suggest that P. pyralis originated in Central- or South America, followed by migration events that populated northern latitudes. Finally, modelling the demographic history of North American P. pyralis serves as a null model of nucleotide diversity patterns, which will inform future studies of adaptation, not only in P. pyralis, but also in other North American taxa.

evolutionary biology↗