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The nature of the molecular and fossil record and their limitations must be ascertained in order to gain the most precise and accurate evolutionary timescale using genetic information. Yet the majority of such timescales are based on point estimates using fossils or the molecular clock. Here we document from the primary literature minimum and maximum fossil age estimates of the divergence of whales from artiodactyls, a commonly used anchor point for calibrating both mitogenomic and nucleogenomic placental timescales.
We applied these reestimates to the most recently established placental timescale based on mitochondrial rRNA and several nuclear loci, and present an attempt to account for both genetic and fossil uncertainty. Our results indicate that disregard for fossil calibration error may inflate the power of the molecular clock when testing the time of ordinal diversification in context with the K-T boundary. However, the early history of placentals, including their superordinal diversification, remained in the Cretaceous despite a conservative approach.
Our conclusions need corroboration across other frequently used fossil anchor points, but also with more genetic partitions on the linear relationship between molecular substitutions and geologic time. Multiple studies of molecular evolutionary timescales agree that the initial diversification of extant placental mammals occurred in the Cretaceous. This result is consistent despite sometimes large variation in molecular divergence time estimation with different calibration points, methods, or datasets Bromham et al.
However, it is often overlooked that substantial error may also exist within a single dataset with a fixed calibration and model of evolution. This error derives from uncertainty in the fossil age of calibration points, branch-length estimation, and tree topology Nei and Kumar In the last 30 years, while phylogenetic and branch-length uncertainties were sometimes acknowledged in error estimation, most molecular clocks have been calibrated with a fixed fossil age.
Fossil records often have been prioritized for estimating molecular rates, specifically what do the branches and nodes in a phylogenetic tree show that are documented in the paleontologic literature as diagnostic e. The tacit assumption of error-free fossil ages from these records and the statistical implications of this assumption have received little attention in the molecular clock literature.
Hence it is unclear what role the specific error associated with empirical data will play in resolution of the placental timescale. Understanding variation in divergence time estimation within and between studies is of particular importance in testing whether placental biodiversity has been marked by major perturbation events across the Cretaceous-Tertiary boundary. Of particular interest is the recent indication that diversification within extant primates, Rodentia, Eulipotyphla, and Xenarthra started in the Late Cretaceous Springer et al.
Nonetheless, because point estimates for the initial diversification within those four orders predate the K-T boundary by only 5—12 million years MYthe effect of molecular error may be especially critical. We investigate the role of genetic and fossil error in resolution of the early placental timescale. The return of mammals to the aquatic realm is considered to be one of the largest macroevolutionary events in amniote history Carroll and is linked with a myriad of distinctive morphological features.
Among mammals, modern whales and dolphins comprise the largest aquatic adaptive radiation combined with the greatest morphological departure from terrestrial ancestors. Because of their distinctive and diagnostic body form, the whale fossil record includes several archaic whales intermediate to modern lineages in time, morphology, and ecology Thewissen Over the last 20 years, important transitional fossils have aided our understanding of how, why, where, and when Cetacea and Artiodactyla diverged Gingerich and Uhen ; Thewissenbut without significantly influencing the time of their initial divergence.
Relative stasis in the timing of cetacean origin has been essential to its utility as a calibration in placental molecular clock studies. In addition, we reviewed molecular clock studies in five journals with frequent focus on mammalian evolution: Molecular Biology and Evolution, Molecular Phylogenetics and Evolution, Journal of Molecular Evolution, Journal of Mammalian Evolutionand Journal of Mammalogy. Our search indicated what is dog food aggression several recent molecular clock studies of placental evolution have favored use of the Artiodactyla-Cetacea divergence to calibrate the remaining mammalian timescale, sometimes in combination with other calibration points.
Those studies used the whale fossil record to estimate the absolute rate of molecular evolution or to force this rate to fall within a range given a rate-heterogeneous molecular dataset. One example is the recent timescale provided by Springer et al. Although six independent fossil records were used to estimate rates across the placental tree, removing the Cetartiodactyl constraint showed the largest shift in divergence time when estimated solely by the remaining five constraints, a sevenfold increase in time estimation error, and yielded younger times for most other nodes.
The removal of the Cetartiodactyl calibration indicates that this constraint may be incongruent with other fossil constraints. Alternatively, evolutionary rate shifts could be especially extensive on the branch leading to Cetartiodactyla, and may not have been corrected for entirely by Bayesian analyses Thorne et al.
Both alternatives merit further investigation. Documentation of the dating methodology what do the branches and nodes in a phylogenetic tree show in calibration fossils or errors associated with molecular clocks is not yet standard procedure. To do so, we must go beyond the often-cited secondary literature. Hence we revisit the primary literature in order to ascertain the details of the earliest Cetartiodactyl fossils and their corresponding stratigraphic horizons.
These details include character descriptions for diagnosis of particular fossils, stratigraphic correlation, and dating methods for the stratigraphically important rocks. From this literature we assess likely minimum and maximum ages for the Artiodactyla-Cetacea divergence and their uncertainties. In this context, our age reestimate of this split is used here to reset the placental timescale using a conservative approach.
Our approach is conservative in its use of minimum fossil age or upper bound Marshall b ; see also Eizirik et al. Within this type of study, erring toward more recent lineage divergence times is considered conservative. The probability of branching at the K-T boundary These analyses are performed on the most what system of linear equation has no solution and largest multilocus available placental dataset Springer et al.
Because difficulty in assigning phylogenetic error to fossils that provide maximal ages impedes statistical resolution, we promote the use of minimum molecular divergence time estimates, following Eizirik et al. We have assumed minimal topologic error for the early history of What do the branches and nodes in a phylogenetic tree show sensu Murphy et al.
Since the Artiodactyla-Cetacea calibration point has been employed in more than 30 molecular studies using different approaches, definitions, and time estimates. A genetics-based estimate of 60 MY no error given for the cow-whale split is commonly used as a calibration point in both mammal and avian mitochondrial genome clocks. Initially estimated at 57 MYA, based on what do the branches and nodes in a phylogenetic tree show 0. Based on these what is a written communication, Arnason et al.
Initially it was uncertain whether the 60 million year estimate of the Artiodactyla-Cetacea ancestor referred to the Cetacea- Hippopotamus or Cetacea-Ruminantia divergence Arnason et al. Use of molecular-based anchor points has been recently criticized Smith what is the best age gap for a relationship Petersonand is ultimately dependent on fossils and not immune to calibration error.
An alternative calibration method using the Artiodactyla-Cetacea divergence is based directly on the Cetartiodactyl fossil record, with a variety of approaches taken. For example, the Artiodactyla-Cetacea divergence has been used in combination with either the minimum age Eizirik et al. These ages ranging between 49 and 63 MY have been applied to both the Cetacea- Hippopotamus node e. The variety and trends observed among users of the Artiodactyla-Cetacea calibration Figure 1 can be traced back to a combination of differing nodal definition, usage of minimum or maximum fossil ages or molecular estimates, and recent finds of older fossils within this lineage Bajpai and Gingerich Molecular data unambiguously support the Hippopotamus as the living artiodactyl sister group of Cetacea e.
This view currently is incongruent with morphological and fossil interpretation, which supports a sister relationship what is a linear relationship equation Cetacea sometimes plus the mesonychid ungulates; O'Leary and Uhen to a monophyletic Artiodactyla Thewissen et al.
However, recent advances have been made toward reconciling both scenarios and indicate that the long-claimed mesonychid-whale grouping is likely due to convergence in dental morphology Naylor and Adams ; Thewissen et al. Another recent advance has been the exciting find of artiodactyl-type anklebones in early hind limb—retaining whales Gingerich et al.
The specific ankle morphology in these early whales is observed in few other groups, including anthracotheres and other early artiodactyls, but excluding mesonychids. Anthracotheres have been considered ancestral to hippos and may provide the fossil link supporting the molecular scenario Gingerich et al. It is possible that cladistic analyses on morphology may not support a whale-hippo grouping because of the highly derived states found in modern and extinct hippos and whales.
Fossils could fill those gaps, but the fossil record of Hippopotamidae traces back to only the Middle Miocene and is too poor on the branch from the Cetacea-Artiodactyl split to document their early evolution O'Leary and Uhen Compared to hippos, can i buy a food lion gift card online earliest history of whales is better understood from fossils of Pakicetidae, arguably the most primitive known whale lineage Thewissen et al.
The oldest fossil directly relevant to the evolution of whales is Himalayacetus subathuensis Figure 2 Bajpai and Gingerich from a marine limestone near the base of the Subathu Formation in the Lesser Himalaya Range, India. Himalayacetus is known from a partial dentition and two molars that show similarities to Pakicetusand hence is placed in Pakicetidae [but see Thewissen and Williams for alternative placement].
Although this fossil is frequently cited as the oldest whale, at 53—54 MYA, its phylogenetic placement and age are somewhat ambiguous. Furthermore, the cetacean synapomorphy, the involucrum of the middle ear Thewissenis not preserved in this fossil. Thus, although likely a whalebased on the teeth and jaw, cladistically Himalayacetus must remain a tentative Cetacean. The material is fragmentary; hence thorough cladistic analysis is difficult O'Leary and Uhen ; Thewissen et al.
The age of Himalayacetus initially was determined from a combination of correlating patterns in local sea level transgression and regression with those described globally, and biochronologically placing the fossil stratum within a benthic foraminifera zone of known age Bajpai and Gingerich According to the same dating techniques and assuming synchrony between the Kuldana and Subathu Formations on the south side of the Himalayas, the previously oldest fossil whales, pakicetids, are younger in age However, the use of benthic, instead of planktonic, foraminifera in biochronological dating is controversial, and the upper Subathu sequence may not have been deposited at the same time as the Pakicetus -bearing Kuldana-Kalakot Formation Thewissen et al.
Nummulites atacicus was used to constrain Himalayacetus to Subathu zones 2—4, but is also found less abundantly in Subathu zones 5—7 Mathuryielding a minimum age of Himalayacetus only slightly older than Pakicetus if synchronous. Mathur later discontinued the use of Nummulites atacicus as a chronological biomarker, and alternative benthic foraminiferan interpretation indicates an age associated with shallow benthic zone 10 Thewissen et al.
The minimum age of Subathu zone 5 corresponds to Biozones are used in biostratigraphy to define zonal limits and refer to biotic events species first and last occurrences. According to molecular analyses, the oldest artiodactyl fossil that can be accurately placed after the split from whales must come from the hippo branch.
If mesonychids belong within Artiodactyla after the hippo-whale split sensu Rosethis would make the oldest mesonychid Dissacusfrom the San Juan Basin in the Torrejonian 2, the oldest fossil within crown Artiodactyla-Cetacea Such an interpretation would yield a 13 million year ghost lineage on the Cetacea branch and implies a 50 million year fossil gap on the hippo branch.
However, considering the new phylogenetic interpretation of mesonychids based on nondental and ankle morphological data Gingerich et al. Thus the age of this divergence is likely better approached by calls wont go through samsung age of the earliest undisputed archaeocetes. The maximum age of the Artiodactyla-Cetacea divergence based on the oldest fossil artiodactyls coincides with the earliest Eocene Diacodexis metsiachus is the oldest artiodactyl bearing an artiodactyl-type ankle Rose Specifically Diacodexis is based on several skeletal fragments, including nearly complete hind limbs, from a paleosol unit about m from the bottom of the Willwood What is the super swipe on bumble What do the branches and nodes in a phylogenetic tree show et al.
This unit can be dated to However, other less complete Diacodexis material has been reported from the earliest Eocene in North America Janis et al. Thus the fossil what is predictor variable in biology of Anthracotheriidae postdates the earliest archaeocetes and cannot be used for estimation of the maximum stem Artiodactyla-Cetacea age.
Molecular data instead support a Paleocene age for crown Artiodactyla Arnason et al. For example, Chriacusan arctocyonid lineage resembling early artiodactyls in its postcranial elements Roseis found in deposits as understanding correlation and causation as Unfortunately, cladistic resolution of the archaic Paleocene mammals is limited and the nature of artiodactyl origin from archaic ungulates remains enigmatic Rose The oldest fossils that display an involucrum in the tympanic region Thewissen provide a minimum age of The first certain fossil cetaceans include the earliest members of Pakicetidae from the redbeds of the lower Kuldana Formation Pakicetus inachusIchthyolestesNalacetus O'Leary and Uhen ; Thewissen et al.
Most of these fossils are based on limited dental material, but recently uncovered postcranial and cranial material Thewissen et al. The lower Kuldana Formation has been considered of Early Eocene Ypresian or earliest Middle Eocene Lutetian age, with the latter placement the most commonly and conservatively accepted age This minimum age is based on radiometric interpolation between the earliest and latest Eocene, and a combination of known magnetostratigraphic reversals linked to foraminifera zonation.
Gingerich and Uhen used What is cause and effect text structure corresponds well with placing the Kuldana Formation at the Wasatchian-Bridgerian boundary Likelihood Gingerich and Uhen and other probabilistic Marshall a statistical methods can be used to put confidence intervals on the age of oldest undisputed fossils, from which a maximum age is inferred based on that known fossil record.
However, an age distribution of several fossil horizons in combination with small stratigraphic ranges is needed to limit the lower bound of the confidence interval, information that often is not available.