Common Lizard Research Articles

Intrinsic climatic predictability affects ornamental coloration of adult males: Evidence for compensation among carotenoid‐ and melanin‐based coloration

Abstract Ornamental coloration is frequently an honest signal of quality associated with sexual selection. While changes in average environmental conditions affect carotenoid‐ and melanin‐based coloration, no evidence exists that changes in intrinsic environmental predictability affect coloration. Here we experimentally manipulated the intrinsic predictability of precipitation in semi‐natural populations of common lizards Zootoca vivipara and tested its effect on ornamental coloration and reproduction of adult males. Less predictable precipitation decreased hue of carotenoid‐based coloration and the extent of melanin‐based coloration. Hue was positively correlated and the extent of melanin‐based coloration was negatively correlated with the male's number of confirmed mate partners. Treatment effects on hue were associated with reduced, and treatment effects on the extent of melanin‐based coloration with increased access to females, in line with differential effects on sexual selection. Neither survival nor reproductive success of adult males significantly differed among predictability treatments, suggesting that negative effects on reproduction caused by changes in carotenoid‐based coloration may have been compensated by changes in melanin‐based coloration. The results also showed that ornamentation, rather than reproduction and survival of adult males, was affected by less favourable conditions. Together with previous results, this suggests that coloration might be more sensitive to changes in environmental conditions than direct fitness proxies. Read the free Plain Language Summary for this article on the Journal blog.

Grandmaternal age at reproduction affects grandoffspring body condition, reproduction and survival in a wild population of lizards

Abstract Age at reproduction can influence the survival and future reproduction of an individual as well as that of their offspring. Remarkably, it has been shown that grandmaternal age at reproduction can also affect the characteristics of grandoffspring in humans and in laboratory or semi‐captive animals. However, currently we do not know whether grandmaternal age effects exist in wild populations. We gathered data on female age at reproduction, offspring and grandoffspring characteristics using a 16‐year long‐term survey of a natural population of the common lizard, Zootoca vivipara. The dataset contains 579 grandoffspring from 135 litters. Body size at birth was not correlated with grandmaternal age at reproduction. However, grandoffspring body condition at birth, grandoffspring survival and reproductive performance of granddaughters were dependent on grandmaternal age. These relationships were independent of maternal age. An age‐structured model showed that the global effect of grandmaternal age was nonlinear and was largely driven by its effect on grandoffspring survival. Fitness was higher for granddaughters produced by grandmothers of intermediate ages. The study shows that age can shape life‐history traits for more than one generation, documenting the importance that grandmaternal age can have in wild populations. A free Plain Language Summary can be found within the Supporting Information of this article.

Understanding the evolution of viviparity using intraspecific variation in reproductive mode and transitional forms of pregnancy.

ABSTRACTHow innovations such as vision, flight and pregnancy evolve is a central question in evolutionary biology. Examination of transitional (intermediate) forms of these traits can help address this question, but these intermediate phenotypes are very rare in extant species. Here we explore the biology and evolution of transitional forms of pregnancy that are midway between the ancestral state of oviparity (egg‐laying) and the derived state, viviparity (live birth). Transitional forms of pregnancy occur in only three vertebrates, all of which are lizard species that also display intraspecific variation in reproductive phenotype. In these lizards (Lerista bougainvillii, Saiphos equalis, and Zootoca vivipara), geographic variation of three reproductive forms occurs within a single species: oviparity, viviparity, and a transitional form of pregnancy. This phenomenon offers the valuable prospect of watching ‘evolution in action’. In these species, it is possible to conduct comparative research using different reproductive forms that are not confounded by speciation, and are of relatively recent origin. We identify major proximate and ultimate questions that can be addressed in these species, and the genetic and genomic tools that can help us understand how transitional forms of pregnancy are produced, despite predicted fitness costs. We argue that these taxa represent an excellent prospect for understanding the major evolutionary shift between egg‐laying and live birth, which is a fundamental innovation in the history of animals.

Resilience of reptiles to megafires.

Extreme climate events, together with anthropogenic land-use changes, have led to the rise of megafires (i.e., fires at the top of the frequency size distribution) in many world regions. Megafires imply that the center of the burnt area is far from the unburnt; therefore, recolonization may be critical for species with low dispersal abilities such as reptiles. We aimed to evaluate the effect of megafires on a reptile community, exploring to what extent reptile responses are spatially shaped by the distance to the unburnt area. We examined the short-term spatiotemporal response of a Mediterranean reptile community after two megafires (>20,000 ha) that occurred in summer 2012 in eastern Spain. Reptiles were sampled over 4 years after the fire in burnt plots located at different distances from the fire perimeter (edge, middle, and center), and in adjacent unburnt plots. Reptile responses were modeled with fire history, as well as climate and remotely sensed environmental variables. In total, we recorded 522 reptiles from 12 species (11 species in the burnt plots and nine in the unburnt plots). Reptile abundance decreased in burnt compared with unburnt plots. The community composition and species richness did not vary either spatially (unburnt and burnt plots) or temporally (during the 4 years). The persistence of reptiles in the burnt area supported their resilience to megafires. The most common lizard species was Psammodromus algirus; both adults and juveniles were found in all unburnt and burnt plots. This species showed lower abundances in burnt areas compared with the unburnt and a slow short-term abundance recovery. The lizard Psammodromus edwarsianus was much less abundant and showed a tendency to increase its abundance in burnt plots compared with unburnt plots. Within the megafire area, P. algirus and P. edwarsianus abundances correlated with the thermal-moisture environment and vegetation recovery regardless of the distance from the fire edge. These results indicated the absence of a short-term reptile recolonization from the unburnt zone, demonstrating that reptiles are resilient (in situ persistence) to megafires when environmental conditions are favorable.

Deduction and exploration of the evolution and function of vertebrate GFPT family.

Glutamine-fructose-6-phosphate aminotransferase (GFPT) is a key factor in the hexosamine metabolism pathway. It regulates the downstream factor O-GlcNAc to change cell function and plays an important role in the metabolism and immune process of tissues and organs. However, the evolutionary relationship of GFPT family proteins in vertebrates has not been elucidated. To deduce and explore the evolution and function of vertebrate GFPT family. 18 GFPT sequences were obtained from Homo sapiens (H. sapiens), Trachypithecus francoisi (T. francoisi), Mus musculus (M. musculus), Rattus norvegicus (R. norvegicus), Gallus gallus (G. gallus), Zootoca vivipara (Z. vivipara), Xenopus tropicalis (X. tropicalis), Danio rerio (D. rerio), Rhincodon typus (R. typus), Plasmodium relictum from National Center for Biotechnology Information (NCBI). The physical and chemical characteristics and molecular evolution of GFPT family proteins and nucleic acid sequences were analyzed by ClustalX2, Gene Doc, MEGA-X, SMART, Datamonkey, R etc. RESULTS: Based on the neighbor-joining (NJ) phylogenetic tree and evolution fingerprints, GFPT family members of vertebrates can be divided into two groups: the GFPT1 group and the GFPT2 group. Seven positive selection sites were identified by IFEL and integrated methods mixed effects model of evolution (MEME) and fixed effects likelihood (REL). Finally, we predicted 28 phosphorylation sites and 18 ubiquitousness sites in the human GFPT1 sequence, 10 phosphorylation sites, and five ubiquitousness sites in GFPT2. Gene ontology (GO) analyzes the protein molecules and KEGG signaling pathways of vertebrates interacting with GFPT family proteins. Our work confirmed that higher animals GFPT family may have differentiated GFPT1 and GFPT2, which meets their own functional needs. This knowledge answers the question what the origin and evolution of GFPT family in vertebrates and provided the basis for disease treatment and function research of GFPT protein.

New record of the viviparous lizard Zootoca vivipara (Jacquin, 1787) in Hungary

The lowland populations of the viviparous lizardZootoca viviparain the Carpathian Basin occur in cold, marshy relict habitats. In one of the largest wetlands in Hungary, Kis-Balaton, in 2016 its presence was confirmed by catching an individual with a small mammal live-trap. This new record is significant, since the nearest known sites of occurrence are at great distance and it is situated between the lowland viviparous populations of the north-northeast and the oviparous populations of the south (in Croatia).

Variation in size and shape sexual dimorphism in the Sceloporus scalaris species group (Squamata: Phrynosomatidae) from the Transvolcanic Belt of Mexico

Abstract We attempted to identify the factors influencing size and shape dimorphism between sexes, as well as among populations and species in the Sceloporus scalaris group (Sceloporus aeneus, S. scalaris, S. bicanthalis and S. subniger). Our analysis focused on five morphological characteristics: snout–vent length, head length, head width, forearm length and tibia length. The effect of environmental variables (precipitation and temperature) on these variables was also tested. We found differences in morphological traits between sexes, and among populations of the same species. The oviparous species (S. aeneus and S. scalaris) were larger in overall body size than the viviparous species (S. bicanthalis and S. subniger). Differences in overall body size among populations were recorded only in S. aeneus and S. scalaris. Male-biased sexual size dimorphism occurred in oviparous but not viviparous lizards (except for one population of S. bicanthalis). An absence of sexual size dimorphism was also recorded in S. subniger and some populations of the remaining species. Two different shape patterns were found; the first was female-biased with larger relative body length in almost all populations, which could be explained by fecundity, and the second was male-biased with relatively larger head and limbs in a few populations, which may be explained by sexual selection. The patterns of sexual size and shape dimorphism show that environment, rather than phylogeny, may be determining the extent of sexual dimorphism. These types of studies show the importance of an integrated evaluation of interpopulation and interspecies variation to determine the factors that generate sexual dimorphism.

Predation on Common Wall Lizards: Survival Probabilities of Melanic Individuals

For estimation of predation plasticine models of prey animals are often used, because the soft material preserves imprints left by predators. We assumed that melanic common wall lizards (Podarcis muralis) disappear by selective predation faster than cryptic individuals and habitat features have important role in this process. We studied the survival probabilities of cryptic and melanic colored plasticine common wall lizard models in habitats with different background coloration on selected places near the city of Pécs (south Hungary), where melanic common wall lizards had been observed earlier. Contrary to our expectations the daily survival rates of melanic plasticine common wall lizards were somewhat higher in all three locations (sandstone quarry, stone wall, coal pit) than those of the cryptic ones, but these differences were not significant. Predators were mostly mammals, which left more marks on plasticine models than birds, but we could not show a preference of the body parts of prey. We concluded that rare occurrence of melanic common wall lizards in habitats near the city of Pécs is not due to predation pressure.

Intraspecific diversity alters the relationship between climate change and parasitism in a polymorphic ectotherm.

Climate-modulated parasitism is driven by a range of factors, yet the spatial and temporal variability of this relationship has received scant attention in wild vertebrate hosts. Moreover, most prior studies overlooked the intraspecific differences across host morphotypes, which impedes a full understanding of the climate-parasitism relationship. In the common lizard (Zootoca vivipara), females exhibit three colour morphs: yellow (Y-females), orange (O-females) and mixed (mixture of yellow and orange, M-females). Zootoca vivipara is also infested with an ectoparasite (Ophionyssus mites). We therefore used this model system to examine the intraspecific response of hosts to parasitism under climate change. We found infestation probability to differ across colour morphs at both spatial (10sites) and temporal (20years) scales: M-females had lower parasite infestations than Y- and O-females at lower temperatures, but became more susceptible to parasites as temperature increased. The advantage of M-females at low temperatures was counterbalanced by their higher mortality rates thereafter, which suggests a morph-dependent trade-off between resistance to parasites and host survival. Furthermore, significant interactions between colour morphs and temperature indicate that the relationship between parasite infestations and climate warming was contingent on host morphotypes. Parasite infestations increased with temperature for most morphs, but displayed morph-specific rates. Finally, infested M-females had higher reductions in survival rates than infested Y- or O-females, which implies a potential loss of intraspecific diversity within populations as parasitism and temperatures rise. Overall, we found parasitism increases with warming temperatures, but this relationship is modulated by host morphotypes and an interaction with temperature. We suggest that epidemiological models incorporate intraspecific diversity within species for better understanding the dynamics of wildlife diseases under climate warming.

The functional genetic architecture of egg-laying and live-bearing reproduction in common lizards.

All amniotes reproduce either by egg-laying (oviparity), which is ancestral to vertebrates or by live-bearing (viviparity), which has evolved many times independently. However, the genetic basis of these parity modes has never been resolved and, consequently, its convergence across evolutionary scales is currently unknown. Here, we leveraged natural hybridizations between oviparous and viviparous common lizards (Zootoca vivipara) to describe the functional genes and genetic architecture of parity mode and its key traits, eggshell and gestation length, and compared our findings across vertebrates. In these lizards, parity trait genes were associated with progesterone-binding functions and enriched for tissue remodelling and immune system pathways. Viviparity involved more genes and complex gene networks than did oviparity. Angiogenesis, vascular endothelial growth and adrenoreceptor pathways were enriched in the viviparous female reproductive tissue, while pathways for transforming growth factor were enriched in the oviparous. Natural selection on these parity mode genes was evident genome-wide. Our comparison to seven independent origins of viviparity in mammals, squamates and fish showed that genes active in pregnancy were related to immunity, tissue remodelling and blood vessel generation. Therefore, our results suggest that pre-established regulatory networks are repeatedly recruited for viviparity and that these are shared at deep evolutionary scales.

Two Novel Adenoviruses in Free-Living British lizards.

In this study, we describe two novel adenoviruses isolated from (i) a common lizard (Zootoca vivipara) found dead and examined post-mortem and (ii) pooled samples from free-living sand lizards (Lacerta agilis agilis). Sequencing indicated the two were closely related atadenovirus strains which were distinct from previously recorded adenoviruses in lizards. Adenoviruses are not always associated with disease in squamates, but morbidity and mortality have been reported. These are the first known cases of adenovirus infection in free-living native British lizards, and further monitoring will be necessary to elucidate the implications of these possible pathogens for vulnerable populations of native reptiles.

Sexual size dimorphism in lizards: Rensch's rule, reproductive mode, clutch size, and line fitting method effects.

Rensch's rule relates to a pattern whereby sexual size dimorphism is more female-biased in small-sized species and more male-biased in large-sized ones. We collected literature and museum data on the body size of males and females belonging to 4032 lizard species, as well as data on their reproductive modes and clutch sizes. We used phylogenetic comparative analyses, and general linear mixed models, to test Rensch's rule and examined how reproductive mode and clutch size affect sexual size dimorphism. Sexual size dimorphism was independent of clutch size in lizard species with variable clutch sizes and in oviparous lizards. Large litters were associated with female-biased sexual dimorphism in viviparous and in scincomorph lizards. Inference regarding Rensch's rule depended on the analytical method used to identify it. The widely used, but less conservative, reduced major axis regression usually support Rensch's rule while ordinary least squares regressions mostly show isometric relationships. The rule tended to apply more to oviparous than to viviparous lizards. We infer that Rensch's rule is, at best, a weak pattern in lizards. This is especially true in viviparous lineages where females reproduce infrequently and therefore evolve large sizes to maximise fecundity, resulting in female-biased dimorphism.

Climatic niche differences among Zootoca vivipara clades with different parity modes: implications for the evolution and maintenance of viviparity

Parity mode (oviparity/viviparity) importantly affects the ecology, morphology, physiology, biogeography and evolution of organisms. The main hypotheses explaining the evolution and maintenance of viviparity are based on bioclimatic predictions and also state that the benefits of viviparity arise during the reproductive period. We identify the main climatic variables discriminating between viviparous and oviparous Eurasian common lizard (Zootoca vivipara) occurrence records during the reproductive period and over the entire year.Analyses based on the climates during the reproductive period show that viviparous clades inhabit sites with less variable temperature and precipitation. On the contrary, analyses based on the annual climates show that viviparous clades inhabit sites with more variable temperatures.Results from models using climates during reproduction are in line with the “selfish-mother hypothesis”, which can explain the success of viviparity, the maintenance of the two reproductive modes, and why viviparous individuals cannot colonize sites inhabited by oviparous ones (and vice versa). They suggest that during the reproductive period viviparity has an adaptive advantage over oviparity in less risky habitats thanks to the selfish behaviour of the mothers. Moreover, the results from both analyses stress that hypotheses about the evolution and maintenance of viviparity need to be tested during the reproductive period.

Global changes explain the long-term demographic trend of the Eurasian common lizard (Squamata: Lacertidae).

The demographic trend of a species depends on the dynamics of its local populations, which can be compromised by local or by global phenomena. However, the relevance of local and global phenomena has rarely been investigated simultaneously. Here, we tested whether local phenomena compromised a species’ demographic trend using the Eurasian common lizard Zootoca vivipara, the terrestrial reptile exhibiting the widest geographic distribution, as a model species. We analyzed the species’ ancient demographic trend using genetic data from its 6 allopatric genetic clades and tested whether its demographic trend mainly depended on single clades or on global phenomena. Zootoca vivipara’s effective population size increased since 2.3 million years ago and started to increase steeply and continuously from 0.531 million years ago. Population growth rate exhibited 2 maxima, both occurring during global climatic changes and important vegetation changes on the northern hemisphere. Effective population size and growth rate were negatively correlated with global surface temperatures, in line with global parameters driving long-term demographic trends. Zootoca vivipara’s ancient demography was neither driven by a single clade, nor by the 2 clades that colonized huge geographic areas after the last glaciation. The low importance of local phenomena, suggests that the experimentally demonstrated high sensitivity of this species to short-term ecological changes is a response in order to cope with short-term and local changes. This suggests that what affected its long-term demographic trend the most, were not these local changes/responses, but rather the important and prolonged global climatic changes and important vegetation changes on the northern hemisphere, including the opening up of the forest by humans.

Население мелких наземных позвоночных животных памятника природы «Чеускинский бор» (Ханты-Мансийский автономный округ - Югра)

There are 25 specially protected natural areas of various status have been organized on the territory of the Khanty-Mansiysk Autonomous Okrug Ugra. Despite their great nature conservation value, the degree of study of flora and fauna, as well as the ecology of plants and animals in the territory of most of them, remains relatively low. Until now, special studies of small terrestrial vertebrates inhabiting the Cheuskinsky Bor have been carried out. In 2020, our research of small terrestrial vertebrates were carried out. The purpose of the study is to identify the species composition and some features of ecology (abundance, biotopic distribution) of small mammals, amphibians and reptiles. The research was carried out in July-September 2020. Metal cones were used to catch most of the vertebrate species. A total of 7400 cone-days have been worked out. 10 species of small mammals and 3 of amphibians are registered. The species composition of small mammals at the Cheuskinsky Bor site is relatively poor, 10 species were captured by cones, 3 were marked visually. The group of dominants included the common shrew and the red-backed vole. They accounted for 22% and 77%, respectively, of all recorded animals. Reptiles in the 2020 season were not registered in the counts, however, according to the literature, the habitat of the common European adder and viviparous lizard is known. Among amphibians, the moor frog and Siberian salamander are rare, the European toad is common. In the future, when conducting annual studies, the list of small mammals can be supplemented by 714 species of insectivores, bats and rodents, the list of amphibians 1 species.

Reptile and Amphibian Diversity and Abundance in an Urban Landscape: Impacts of Fragmentation and the Conservation Value of Small Patches

Urbanization is a major contributor to habitat loss and fragmentation and is considered a global threat to biodiversity. We studied reptile and amphibian species diversity and abundance in a highly fragmented landscape adjacent to the second largest metropolitan area in the United States. Habitat patches in our study area were made up of remnant native vegetation surrounded by roads, housing, and other urban development. Species richness and diversity were positively associated with patch size, but patch age was not significantly associated with community characteristics. Four relatively common species were not detected in the small patches, indicating the possibility they had been extirpated by the time monitoring began, and six rarer species were not detected or detected only once in these patches. Although the patch size effect on species diversity was strong, we found that several of the small habitat patches had similar diversity to large patches, indicating potential value of these small habitat patches in protecting species as “microreserves.” In addition, one lizard species was found to be significantly more abundant in the smaller patches. To determine if abundance changed over time, we compared capture rates for four common lizards at the same sites ten years later. For three of the four species, abundance decreased over that period, specifically in the small patches. Although our long-term monitoring has confirmed that the full suite of herpetofauna is currently preserved in the study area overall, declines even in the common species over time hint at the potential severity of the threat of urbanization to rare species.

Individual telomere dynamics and their links to life history in a viviparous lizard

Emerging patterns suggest telomere dynamics and life history are fundamentally linked in endotherms through life-history traits that mediate the processes underlying telomere attrition. Unlike endotherms, ectotherms maintain the ability to lengthen somatic telomeres throughout life and the link between life-history strategies and ectotherm telomere dynamics is unknown. In a well-characterized model system (Niveoscincus ocellatus), we used long-term longitudinal data to study telomere dynamics across climatically divergent populations. We found longer telomeres in individuals from the cool highlands than those from the warm lowlands at birth and as adults. The key determinant of adult telomere length across populations was telomere length at birth, with population-specific effects of age and growth on adult telomere length. The reproductive effort had no proximate effect on telomere length in either population. Maternal factors influenced telomere length at birth in the warm lowlands but not the cool highlands. Our results demonstrate that life-history traits can have pervasive and context-dependent effects on telomere dynamics in ectotherms both within and between populations. We argue that these telomere dynamics may reflect the populations' different life histories, with the slow-growing cool highland population investing more into telomere lengthening compared to the earlier-maturing warm lowland population.

Intense nocturnal warming alters growth strategies, colouration and parasite load in a diurnal lizard.

In the past decades, nocturnal temperatures have been playing a disproportionate role in the global warming of the planet. Yet, they remain a neglected factor in studies assessing the impact of global warming on natural populations. Here, we question whether an intense augmentation of nocturnal temperatures is beneficial or deleterious to ectotherms. Physiological performance is influenced by thermal conditions in ectotherms and an increase in temperature by only 2°C is sufficient to induce a disproportionate increase in metabolic expenditure. Warmer nights may expand ectotherms' species thermal niche and open new opportunities for prolonged activities and improve foraging efficiency. However, increased activity may also have deleterious effects on energy balance if exposure to warmer nights reduces resting periods and elevates resting metabolic rate. We assessed whether warmer nights affected an individual's growth, dorsal skin colouration, thermoregulation behaviour, oxidative stress status and parasite load by exposing yearling common lizards (Zootoca vivipara) from four populations to either ambient or high nocturnal temperatures for approximately 5weeks. Warmer nocturnal temperatures increased the prevalence of ectoparasitic infestation and altered allocation of resources towards structural growth rather than storage. We found no change in markers for oxidative stress. The thermal treatment did not influence thermal preferences, but influenced dorsal skin brightness and luminance, in line with a predicted acclimation response in colder environments to enhance heat gain from solar radiation. Altogether, our results highlight the importance of considering nocturnal warming as an independent factor affecting ectotherms' life history in the context of global climate change. ​.

The role of social costs as a mechanism enforcing the honesty of ultraviolet-reflecting signals in a lizard

AbstractAccording to animal signalling theory, social costs incurred by aggressive conspecifics are one mechanism maintaining signal honesty. Although our understanding of signal evolution has much improved for pigment-based colours, the mechanisms maintaining the honesty of structural colour signals, such as ultraviolet (UV), remain elusive. Here, we used the common lizard (Zootoca vivipara) to test whether the honesty of UV-reflecting signals displayed on male throats is under social control. To do so, we staged agonistic interactions between non-manipulated focal males and opponents of either larger or smaller body size. We manipulated the UV component of the male throat colour patch to create small cheaters with UV-enhanced throats, large cheaters with UV-reduced throats, and their respective controls. In support of a conventional signal hypothesis, focal males were aggressive towards large cheaters and became submissive when these large cheaters retaliated, and were less submissive against small cheaters. However, that focal males were not more aggressive towards small cheaters contradicts our initial predictions. We confirm that male UV reflectance and bite force were good predictors of contest outcomes in control conditions. Overall, we provide partial evidence suggesting that social costs enforce UV signal honesty in common lizards.

The Allelic Variants in Microsatellite Loci and Phylogenetic Relationships of <i>Darevskia rudis</i> (Bedriaga, 1886) and <i>D. bithynica</i> (Méhely, 1909) Based on Mitochondrial DNA in Turkey

The spiny-tailed lizard, which has a series of taxonomic revisions, is one of the most common lizard species in Turkey. In this study, sequence data derived from three microsatellite loci (Du215, Du281, and Du323), two mitochondrial (16S rRNA and Cyt-b) genes and combined data were used to evaluate the taxonomic status of Darevskia rudis and Darevskia bithynica with new samples from all subspecies populations in Turkey. Our results indicated that the genetic variations of microsatellite loci were not specific to populations within species, and only minor differences separated D. rudis and D. bithynica populations. Furthermore, the markers we used for phylogenetic analyses (NJ, ML, MP, and BI) produced topologically similar trees based on 16S rRNA and Cyt-b while the combined data produced conflicting trees with the separate gene analyses. Finally, the basal relationships among the populations in Turkish populations D. rudis and D. bithynica were not resolved with this dataset, and we found a hard polytomy at the basis of the phylogeny.