Middle Eocene rodents from Peruvian Amazonia reveal the pattern and timing of caviomorph origins and biogeography

The long-term isolation of South America during most of the Cenozoic produced a highly peculiar terrestrial vertebrate biota, with a wide array of mammal groups, among which caviomorph rodents and platyrrhine primates are Mid-Cenozoic immigrants. In the absence of indisputable pre-Oligocene South American rodents or primates, the mode, timing and biogeography of these extraordinary dispersals remained debated. Here, we describe South America's oldest known rodents, based on a new diverse caviomorph assemblage from the late Middle Eocene (approx. 41 Ma) of Peru, including five small rodents with three stem caviomorphs. Instead of being tied to the Eocene/Oligocene global cooling and drying episode (approx. 34 Ma), as previously considered, the arrival of caviomorphs and their initial radiation in South America probably occurred under much warmer and wetter conditions, around the Mid-Eocene Climatic Optimum. Our phylogenetic results reaffirm the African origin of South American rodents and support a trans-Atlantic dispersal of these mammals during Middle Eocene times. This discovery further extends the gap (approx. 15 Myr) between first appearances of rodents and primates in South America.     

Main radiation events in Pan‐Octodontoidea (Rodentia,Caviomorpha)

Caviomorphs (South American hystricognaths) are recorded in the continent since the middle Eocene. The middle Eocene–early Oligocene is considered a key moment for their evolutionary history because by the early Oligocene they were differentiated into four superfamilies: Octodontoidea, Cavioidea, Chinchilloidea and Erethizontoidea. Due to their generalized dental patterns and abundance in the fossil record, Octodontoidea are interesting for analysing the origin and early history of caviomorphs. The phylogenetic relationships of the earliest octodontoids are studied herein. Results confirmed a basal caviomorph diversification in the middle Eocene (c. 45 Mya), with one lineage leading to Pan‐Octodontoidea, and another leading to Erethizontoidea, Cavioidea and Chinchilloidea, which is not in accordance with analyses based on molecular data. Three major radiations were identified: the first one (late Eocene?/early Oligocene?) occurred in low latitudes with the differentiation of Pan‐Octodontoidea and the earliest crown‐Octodontoidea. The second radiation (late Oligocene) was a large‐scale South American event; in the southernmost part of the continent it is recognized as the first Patagonian octodontoid radiation, which provided the characteristic high morphological disparity of the superfamily. The third radiation (late Miocene) is characterized by the replacement of ‘old’ by ‘modern’ octodontoids; the nature of this third event needs to be study in a broader taxonomic context. © 2015 The Linnean Society of London     

A newly recognized family from the Horn of Africa,the Heterocephalidae (Rodentia: Ctenohystrica)

The Ctenohystrica is one of the three major lineages of rodents and contains diverse forms related to gundis, porcupines, and guinea pigs. Phylogenetic analyses of this group using mitochondrial and nuclear gene sequences confirm the monophyly of the infraorder Hystricognathi and most of its recognized subclades, including both the Neotropical caviomorphs and the African phiomorphs, which are recovered as sister groups. Molecular timetrees calibrated with 22 securely placed fossils indicate that hystricognath superfamilies originated in the Eocene and Oligocene and most families had appeared by the end of the Oligocene, ～23 Mya. Divergences leading to hystricognath genera took place in the Miocene and Pliocene, with a single exception. The naked mole‐rat (Heterocephalus) diverged from other African mole‐rats (Bathyergidae) in the early Oligocene (～31.2 Mya), when the four caviomorph superfamilies (Erethizonoidea and Cavioidea at 32.4 Mya, Chinchilloidea and Octodontoidea at 32.8 Mya) were first appearing in South America. The extended independent evolution of Heterocephalus suggested by this analysis prompted a closer examination of mole‐rat characters. Heterocephalus indeed shares many characters with bathyergids, befitting their joint membership in the parvorder Bathyergomorphi and superfamily Bathyergoidea as well as their shared exploitation of subterranean lifestyles. However, a diverse array of cranial, dental, postcranial, external, and ecological characters distinguishes Heterocephalus from other African mole‐rats. These differences equal or exceed those used to diagnose caviomorph families and justify recognizing the naked mole‐rat in its own family, Heterocephalidae Landry, 1957. This taxonomic arrangement poses questions for the inter‐relationships of fossil and extant mole‐rats and brings time equivalence to the ranks assigned to the major clades of hystricognaths. © 2014 The Linnean Society of London     

Asynchronous colonization of Madagascar by the four endemic clades of primates, tenrecs, carnivores, and rodents as inferred from nuclear genes

Madagascar harbors four large adaptive radiations of endemic terrestrial mammals: lemurs, tenrecs, carnivorans, and rodents. These rank among the most spectacular examples of evolutionary diversification, but their monophyly and origins are debated. The lack of Tertiary fossils from Madagascar leaves molecular studies as most promising to solve these controversies. We provide a simultaneous reconstruction of phylogeny and age of the four radiations based on a 3.5-kb data set from three nuclear genes (ADRA2B, vWF, and AR). The analysis supports each as a monophyletic clade, sister to African taxa, and thereby identifies four events of colonization out of Africa. To infer the time windows for colonization, we take into account both the divergence from the closest non-insular sister group and the initial intra-insular radiation, which is a novel but conservative approach in studies of the colonization history of Madagascar. We estimate that lemurs colonized Madagascar between 60 million years ago (Mya) (split from lorises) and 50 Mya (lemur radiation) (70-41 Mya taking 95% credibility intervals into account), tenrecs between 42 and 25 Mya (50-20 Mya), carnivorans between 26 and 19 Mya (33-14 Mya), and rodents between 24 and 20 Mya (30-15 Mya). These datings suggest at least two asynchronous colonization events: by lemurs in the Late Cretaceous-Middle Eocene, and by carnivorans and rodents in the Early Oligocene-Early Miocene. The colonization by tenrecs may have taken place simultaneously with either of these two events, or in a third event in the Late Eocene-Oligocene. Colonization by at least lemurs, rodents, and carnivorans appears to have occurred by overseas rafting rather than via a land bridge hypothesized to have existed between 45 and 26 Mya, but the second scenario cannot be ruled out if credibility intervals are taken into account.     

Extinction of South American sparassodontans (Metatheria): environmental fluctuations or complex ecological processes?

Sparassodontans are a diverse but now extinct group of metatherians that were apex predators in South America during most of the Cenozoic. Studying their decline has been controversial mainly due to the scarcity of the fossil record, and different methodological approaches have led to contradictory hypotheses. In an effort to explore questions about their extinction, we developed a novel multi‐model statistical approach to analyse all of the currently available data at a continental scale. Using multiple regression analysis and new advances in beta diversity analysis, we used all currently available fossil data at a continental scale to test four competing hypotheses to account for the decline of sparassodontans: competition with placental carnivorans, competition with avian phorusrhacids, non‐competitive ecological interactions, and environmental fluctuations. Our results show that the sparassodontan extinction was a gradual process with species disappearing throughout the Cenozoic. Multiple regression analysis supported non‐competitive ecological interactions as the best extinction model. Native South American ungulates, African migrants (caviomorph rodents and platyrrhine primates) and didelphimorphians were the groups with the highest statistical significance. Sparassodontan beta diversity increased between South American Land Mammal Ages after the Paleocene–Eocene boundary. Our results demonstrate that ecological modelling techniques illuminate aspects of extinction processes whilst mitigating the limitations of the fossil record. Our study suggests that non‐competitive ecological interactions could have been the main driver for sparassodontan extinction rather than, as commonly assumed, a result of competition and/or abiotic fluctuations.     

Phylogeny and biogeography of the carnivorous plant family Sarraceniaceae

The carnivorous plant family Sarraceniaceae comprises three genera of wetland-inhabiting pitcher plants: Darlingtonia in the northwestern United States, Sarracenia in eastern North America, and Heliamphora in northern South America. Hypotheses concerning the biogeographic history leading to this unusual disjunct distribution are controversial, in part because genus- and species-level phylogenies have not been clearly resolved. Here, we present a robust, species-rich phylogeny of Sarraceniaceae based on seven mitochondrial, nuclear, and plastid loci, which we use to illuminate this family's phylogenetic and biogeographic history. The family and genera are monophyletic: Darlingtonia is sister to a clade consisting of Heliamphora+Sarracenia. Within Sarracenia, two clades were strongly supported: one consisting of S. purpurea, its subspecies, and S. rosea; the other consisting of nine species endemic to the southeastern United States. Divergence time estimates revealed that stem group Sarraceniaceae likely originated in South America 44-53 million years ago (Mya) (highest posterior density [HPD] estimate = 47 Mya). By 25-44 (HPD = 35) Mya, crown-group Sarraceniaceae appears to have been widespread across North and South America, and Darlingtonia (western North America) had diverged from Heliamphora+Sarracenia (eastern North America+South America). This disjunction and apparent range contraction is consistent with late Eocene cooling and aridification, which may have severed the continuity of Sarraceniaceae across much of North America. Sarracenia and Heliamphora subsequently diverged in the late Oligocene, 14-32 (HPD = 23) Mya, perhaps when direct overland continuity between North and South America became reduced. Initial diversification of South American Heliamphora began at least 8 Mya, but diversification of Sarracenia was more recent (2-7, HPD = 4 Mya); the bulk of southeastern United States Sarracenia originated co-incident with Pleistocene glaciation, <3 Mya. Overall, these results suggest climatic change at different temporal and spatial scales in part shaped the distribution and diversity of this carnivorous plant clade.     

Major Radiations in the Evolution of Caviid Rodents: Reconciling Fossils,Ghost Lineages,and Relaxed Molecular Clocks

Background

Caviidae is a diverse group of caviomorph rodents that is broadly distributed in South America and is divided into three highly divergent extant lineages: Caviinae (cavies), Dolichotinae (maras), and Hydrochoerinae (capybaras). The fossil record of Caviidae is only abundant and diverse since the late Miocene. Caviids belongs to Cavioidea sensu stricto (Cavioidea s.s.) that also includes a diverse assemblage of extinct taxa recorded from the late Oligocene to the middle Miocene of South America (“eocardiids”).

Results

A phylogenetic analysis combining morphological and molecular data is presented here, evaluating the time of diversification of selected nodes based on the calibration of phylogenetic trees with fossil taxa and the use of relaxed molecular clocks. This analysis reveals three major phases of diversification in the evolutionary history of Cavioidea s.s. The first two phases involve two successive radiations of extinct lineages that occurred during the late Oligocene and the early Miocene. The third phase consists of the diversification of Caviidae. The initial split of caviids is dated as middle Miocene by the fossil record. This date falls within the 95% higher probability distribution estimated by the relaxed Bayesian molecular clock, although the mean age estimate ages are 3.5 to 7 Myr older. The initial split of caviids is followed by an obscure period of poor fossil record (refered here as the Mayoan gap) and then by the appearance of highly differentiated modern lineages of caviids, which evidentially occurred at the late Miocene as indicated by both the fossil record and molecular clock estimates.

Conclusions

The integrated approach used here allowed us identifying the agreements and discrepancies of the fossil record and molecular clock estimates on the timing of the major events in cavioid evolution, revealing evolutionary patterns that would not have been possible to gather using only molecular or paleontological data alone.     

The role of Asia in the origin and diversification of hystricognathous rodents

In the absence of a comprehensive pre‐Oligocene fossil record, the origin and early evolution of hystricognathous rodents have long been the subject of much uncertainty. Baluchimyinae (Rodentia) were initially interpreted as a subfamily of the ctenodactyloid Chappatimyidae (sciurognathous), a group considered to be endemic to the Indian subcontinent and to be closely related to hystricognathous rodents. A newly discovered early Oligocene hystricognathous rodent, Bugtimys zafarullahi gen. n. et sp. n., described herein, from the Bugti Hills (Balochistan, Pakistan) sheds new light on the higher level taxonomy of the previously described Baluchimyinae. As a contribution to the phylogenetic debates regarding the origin of Hystricognathi, we present a cladistic assessment of the dental evidence for the Palaeogene hystricognathous rodent cladogenesis. Our phylogenetic results consistently support the monophyly of the Hystricognathiformes clade (including Tsaganomyidae plus Hystricognathi) of which baluchimyine rodents are clearly members. There is, however, no support for the monophyly of a baluchimyine clade. Nonetheless, ‘baluchimyines’ are for the moment reinterpreted as Hystricognathi incertae sedis. Hystricognathous rodents appear to be well diversified at least since the early Oligocene, both in Africa and South America (phiomorphs and caviomorphs, respectively), and also now in south Asia. Furthermore, our phylogenetic results support close relationships between early hystricognathous and Asian ‘ctenodactyloid’ rodents, which clearly points to an Asian origin for Hystricognathi. In this phylogenetic framework, ‘baluchimyines’ and tsaganomyids are representatives of an initial phase of diversification of hystricognathous rodents in Asia. Oligocene phiomorphs and caviomorphs (sister groups) seem therefore to share a common ‘Asian’ hystricognathous ancestor. This reinforces the possibility that the early dispersal of hystricognathous rodents to South America was not from Africa but from Asia.     

Conservation of placentation during the tertiary radiation of mammals in South America

The eutherian placenta is considered to possess great plasticity, but it is not clear how this variation reflects adaptation to different ecological niches. Because South America was isolated for most of the Tertiary, it represents a natural laboratory to examine this question. We here describe placentation in three South American groups: Xenarthra have been part of the fauna from at least the mid‐Paleocene whereas caviomorph rodents and Neotropical primates are each derived from a single founder that reached South America in the Eocene and Oligocene, respectively. The common ancestor of Xenarthra had a villous, haemochorial placenta, from which the labyrinthine, endotheliochorial placenta of sloths later evolved. Placentation in Caviomorpha follows an extraordinary stable pattern, characterized by a haemomonochorial, labyrinthine and highly lobed structure with specialized growing areas. This pattern was present before arrival of these rodents in South America and enabled a successful radiation especially during the spread of grasslands. Neotropical primates have haemochorial, trabecular placentas with a specialized maternal blood supply; a pattern that contrasts with that of Old World monkeys and may have been present in the founder generation on arrival in South America. In conclusion, there is a dichotomy within Xenarthra but otherwise the ancient South American mammals do not show much variation in principal placental characters. Thus, the successful radiation of these three groups, and their adaptation to diverse ecological niches, did not require substantial alterations in placentation. J. Morphol., 2013. © 2013 Wiley Periodicals, Inc.     

'First' appearances in the Cenozoic land-mammal record of the Greater Antilles: significance and comparison with South American and Antarctic records

Aim Through analysis of fossil records, the aim of this paper is to show that fossil representatives of at least three land‐mammal clades (pitheciine atelid primates, heteropsomyine echimyid rodents, and megalonychid phyllophagan xenarthrans) that once lived in the Greater Antilles are as old as, if not older than, ‘first’ occurrences of these same groups on the South American mainland. Location Greater Antilles, South America, Antarctic Peninsula. Methods Analysis of Cenozoic land‐mammal fossil records for the three areas. Results Comparison reveals an interesting similarity to the Tertiary vertebrate palaeontological record for the Antarctic Peninsula (Seymour Island), in the sense that the latter also includes early (Eocene) representatives of some typical ‘South American’ groups (e.g. meridiungulates, sloths, certain marsupial groups). Conclusions Given how limited the Antillean and Antarctic records are in quantity and quality, it seems unlikely that these ‘first’ appearances have much bearing on real origins (basal divergences). Rather, it suggests that the fossil basis for interpreting the origin and earliest diversification of ‘South American’ clades during the latest Cretaceous/early Cenozoic is probably even scantier than generally realized. In particular, the Antillean record strengthens arguments that some crown‐group continental lineages are considerably older than fossil evidence currently allows – a point increasingly (if unevenly) supported by molecular studies of many of the same clades.     

Molecular estimation of eulipotyphlan divergence times and the evolution of "Insectivora"

"Insectivores" are one of the key groups in understanding mammalian origins. For years, systematics of "Lipotyphla" taxa remained extremely unstable and challenged. Today, with the application of molecular techniques, "Lipotyphla" appears to be a paraphyletic assemblage that encompasses hedgehogs, shrews, and moles (i.e., Eulipotyphla-a member of Laurasiatheria), and golden moles and tenrecs (i.e., Afrosoricida-a member of Afrotheria). Based on nuclear genes and on this well-established phylogenetic framework, we estimated Bayesian relaxed molecular clock divergence times among major lineages of "Lipotyphla." Crown placental mammals are shown to diversify 102+/-6 million years ago (Mya; mean+/-one standard-deviation), followed by Boreoeutheria (94+/-6 Mya), Laurasiatheria (85+/-5 Mya), and Eulipotyphla (73+/-5), with moles separating from hedgehogs+shrews just at the K/T boundary (65+/-5 Mya). During the Early and Middle Eocene, all extant eulipotyphlan subfamilies originated: Uropsilinae (52+/-5 Mya), and Desmaninae, Talpinae, Erinaceinae, Hylomyinae, Soricinae, and Crocidurinae (38-42+/-5 Mya). Afrosoricida separated from Macroscelidae 69+/-5 Mya, golden moles from tenrecs 63+/-5 Mya, and the diversification within tenrecs occurred 43+/-5 Mya. Divergence times are shown to be in reasonably good agreement with the fossil record of eulipotyphlans, but not with the one of afrosoricid "insectivores." Eulipotyphlans diversification might have been sculpted by variations in paleoclimates of the cenozoic era.     

Intraspecific variation in space use, group size, and mating systems of caviomorph rodents

Intraspecific variation in social systems is widely recognized across many taxa, and specific models, including polygamy potential, resource defense, and resource dispersion, have been developed to explain the relationship between ecological variation and social organization. Although mammals from temperate North America and Eurasia have provided many insights into this relationship, rodents from the Neotropics and temperate South America have largely been ignored. In this review we focus on reports documenting intraspecific variation in spacing systems, group size, and mating systems of caviomorphs. This large group of New World hystricognath rodents occupies a diverse array of habitats; thus, members of the same species potentially exhibit different social systems in response to different ecological conditions. Spatial patterns vary in response to a diverse array of factors, including predation, food availability, population density, and soil characteristics. Changes in group size typically correlate with changes in resource availability, particularly food. Mating systems generally reflect the ability of males to control access to females, which may depend on population density or food distribution. In general, social organization in caviomorphs fits predictions of resource-based models; however, most studies have been purely observational, involving small numbers of animals over short time periods and reporting qualitative rather than quantitative levels of ecological correlates. In future studies the use of molecular techniques and controlled, experimental manipulations can increase our understanding of intraspecific variation in caviomorph social systems. This understudied group of rodents offers excellent opportunities to provide insights into the influence of ecological conditions on behavior such as social systems.     

Primate molecular divergence dates

With genomic data, alignments can be assembled that greatly increase the number of informative sites for analysis of molecular divergence dates. Here, we present an estimate of the molecular divergence dates for all of the major primate groups. These date estimates are based on a Bayesian analysis of approximately 59.8 kbp of genomic data from 13 primates and 6 mammalian outgroups, using a range of paleontologically supported calibration estimates. Results support a Cretaceous last common ancestor of extant primates (approximately 77 mya), an Eocene divergence between platyrrhine and catarrhine primates (approximately 43 mya), an Oligocene origin of apes and Old World monkeys (approximately 31 mya), and an early Miocene (approximately 18 mya) divergence of Asian and African great apes. These dates are examined in the context of other molecular clock studies.     

Molecular dating: ape bones agree with chicken entrails

Molecular time estimates, especially those that employed the 310 million years ago (Mya) date of mammal-bird divergence as the calibration point, were criticized in recent publications. In this article, we estimate the divergence time of primates and rodents, primates and artiodactyls and the different great ape species by using two independent calibration-time ranges and maximally conservative error estimates. We observed a variation of approximately +/-15-20% for most of the molecular time estimates in the 10-100 Mya range. The estimated range of the primate-rodent divergence time, 84-121 Mya, includes the date obtained with the 310 million years calibration point (110 Mya). We conclude that molecular time estimates remain useful tools of evolutionary biology, although utmost caution is required when interpreting the results.     

Recent advances in South American mammalian paleontology

Recently discovered deposits containing terrestrial mammal fossils, together with multidisciplinary studies of classical sequences, have yielded dramatic insights into the biotic and environmental history of South America. Notable advances include several new fossil primate taxa, an improved chronology of two major immigration events (caviomorph rodents and new world monkeys), documentation of the oldest mammalian faunas dominated by grazing taxa (which suggests that grasslands appeared at least 15 million years earlier than on other continents), evidence of early biogeographical provinciality within South America, and improved sampling of the best known Cenozoic tropical South American paleofauna.     

Eocene Diversification of Crown Group Rails (Aves: Gruiformes: Rallidae)

Central to our understanding of the timing of bird evolution is debate about an apparent conflict between fossil and molecular data. A deep age for higher level taxa within Neoaves is evident from molecular analyses but much remains to be learned about the age of diversification in modern bird families and their evolutionary ecology. In order to better understand the timing and pattern of diversification within the family Rallidae we used a relaxed molecular clock, fossil calibrations, and complete mitochondrial genomes from a range of rallid species analysed in a Bayesian framework. The estimated time of origin of Rallidae is Eocene, about 40.5 Mya, with evidence of intrafamiliar diversification from the Late Eocene to the Miocene. This timing is older than previously suggested for crown group Rallidae, but fossil calibrations, extent of taxon sampling and substantial sequence data give it credence. We note that fossils of Eocene age tentatively assigned to Rallidae are consistent with our findings. Compared to available studies of other bird lineages, the rail clade is old and supports an inference of deep ancestry of ground-dwelling habits among Neoaves.     

Craniodental morphology and systematics of a new family of hystricognathous rodents (Gaudeamuridae) from the late eocene and early oligocene of Egypt

Background

Gaudeamus is an enigmatic hystricognathous rodent that was, until recently, known solely from fragmentary material from early Oligocene sites in Egypt, Oman, and Libya. Gaudeamus'' molars are similar to those of the extant cane rat Thryonomys, and multiple authorities have aligned Gaudeamus with Thryonomys to the exclusion of other living and extinct African hystricognaths; recent phylogenetic analyses have, however, also suggested affinities with South American caviomorphs or Old World porcupines (Hystricidae).

Methodology/Principal Findings

Here we describe the oldest known remains of Gaudeamus, including largely complete but crushed crania and complete upper and lower dentitions. Unlike younger Gaudeamus species, the primitive species described here have relatively complex occlusal patterns, and retain a number of plesiomorphic features. Unconstrained parsimony analysis nests Gaudeamus and Hystrix within the South American caviomorph radiation, implying what we consider to be an implausible back-dispersal across the Atlantic Ocean to account for Gaudeamus'' presence in the late Eocene of Africa. An analysis that was constrained to recover the biogeographically more plausible hypothesis of caviomorph monophyly does not place Gaudeamus as a stem caviomorph, but rather as a sister taxon of hystricids.

Conclusions/Significance

We place Gaudeamus species in a new family, Gaudeamuridae, and consider it likely that the group originated, diversified, and then went extinct over a geologically brief period of time during the latest Eocene and early Oligocene in Afro-Arabia. Gaudeamurids are the only known crown hystricognaths from Afro-Arabia that are likely to be aligned with non-phiomorph members of that clade, and as such provide additional support for an Afro-Arabian origin of advanced stem and basal crown members of Hystricognathi.     

INVESTIGATING PROCESSES OF NEOTROPICAL RAIN FOREST TREE DIVERSIFICATION BY EXAMINING THE EVOLUTION AND HISTORICAL BIOGEOGRAPHY OF THE PROTIEAE (BURSERACEAE)

Andean uplift and the collision of North and South America are thought to have major implications for the diversification of the Neotropical biota. However, few studies have investigated how these geological events may have influenced diversification. We present a multilocus phylogeny of 102 Protieae taxa (73% of published species), sampled pantropically, to test hypotheses about the relative importance of dispersal, vicariance, habitat specialization, and biotic factors in the diversification of this ecologically dominant tribe of Neotropical trees. Bayesian fossil‐calibrated analyses date the Protieae stem at 55 Mya. Biogeographic analyses reconstruct an initial late Oligocene/early Miocene radiation in Amazonia for Neotropical Protieae, with several subsequent late Miocene dispersal events to Central America, the Caribbean, Brazil's Atlantic Forest, and the Chocó. Regional phylogenetic structure results indicate frequent dispersal among regions throughout the Miocene and many instances of more recent regional in situ speciation. Habitat specialization to white sand or flooded soils was common, especially in Amazonia. There was one significant increase in diversification rate coincident with colonization of the Neotropics, followed by a gradual decrease consistent with models of diversity‐dependent cladogenesis. Dispersal, biotic interactions, and habitat specialization are thus hypothesized to be the most important processes underlying the diversification of the Protieae.     

广西南宁渐新世基于咽齿的鲤亚科一新属

记述了产自广西南宁盆地渐新世早—中期的鲤科鲤亚科一新属种——伍氏南宁鲤(+Nanningocyprinus wui gen.et sp.nov).化石材料包括一些咽喉骨和咽齿.这些咽喉骨和咽齿的以下特征组合明显不同于其他鲤科鱼类:齿式-3·2·1,主行第一枚咽齿大小是第二枚的3～4倍,第二枚咽齿及第二行第一枚咽齿咀嚼面上各仅有一条沟纹,咽骨前角很发育.该属种的发现进一步证明鲤亚科是鲤科中较早的分支,中国南方很有可能是鲤亚科的起源和分化中心.