Climatic and biogeographical associations of southern African dung beetles (Coleoptera: Scarabaeidae s. str.)

Climatic and biogeographical associations of southern African dung beetles (Scarabaeinae, Coprinae) were analysed from a collection amassed between 1971 and 1986. Endemism to Africa south of 15°S was much greater in southwesterly climates (winter rainfall, bimodal spring/autumn rainfall, arid late summer rainfall) than to the north‐east in the moist, mid‐summer rainfall region. Major biogeographical groups centred to the south‐west comprised predominantly southern African endemics, Western/Eastern Cape coast, Karoo, Karoo/Namib, northern Namibia and the south‐western Kalahari. Biogeographical groups centred on the south‐eastern highlands and the subtropical east coast (mid‐summer rainfall) also comprised predominantly southern African endemics. All other major groups centred to the north‐east in the mid‐summer rainfall region comprised predominantly species with widespread tropical biogeographical affiliations, pan‐southern Africa but centred in the east, pan‐mid‐summer rainfall region, eastern mid‐summer rainfall region, tropical east Zimbabwe/central Moçambique, subtropical/tropical game reserves (non‐ruminant dung specialists). There were cross‐climatic differences in taxonomic composition of the fauna. Within the winter rainfall region, percentage species composition of Scarabaeinae was greater whereas that of the coprine tribe, Onthophagini, was lower than within the other three climatic regions. Percentage species richness of most other tribes of Coprinae differed little between most climates but that of Scarabaeinae declined from west to east (Canthonini, Scarabaeini), east to west (Sisyphini) or to either side of the late summer rainfall region (Gymnopleurini).     

Dung feeding in adult scarabaeines (tunnellers and endocoprids): even large dung beetles eat small particles

Abstract 1. The maximum size of ingested particles was determined in 15 species of adult dung beetle (Scarabaeidae: Scarabaeinae) by mixing small latex or glass balls of known diameter into the dung used as food. Twelve species (tribes Coprini, Onitini, Oniticellini, and Onthophagini) were tunnellers (making dung stores for feeding and breeding in the soil below the pat) and three species (tribe Oniticellini) were endocoprids (feeding and breeding in the dung pat itself).
2. The test species, covering a wide range of body size (fresh weights 0.05–7.4 g), ingested minute particles only (maximum diameter 8–50 µm), and there was a statistically significant but numerically small increase in particle size with body weight.
3. When the effect of body size was taken into account, taxon (tribe), ecological group (tunneller/endocoprid), and dung preference (coarse/fine) had no significant effect on the size of ingested particles.
4. Tests using two tunnelling species did not indicate that beetles use their mandibles to grind dung particles prior to ingestion.
5. The results suggest essentially the same feeding mechanism in all adult tunnelling or endocoprid scarabaeines that eat fresh dung. Larger, indigestible plant fragments are avoided by filtration, and ingestion is confined to very small particles of higher nutritional value.     

Mitogenome of Coprophanaeus ensifer and phylogenetic analysis of the Scarabaeidae family (Coleoptera)

Several studies about the phylogenetic relationships of the Scarabaeinae subfamily (Coleoptera: Scarabaeidae) have been performed, but some phylogenetic uncertainties persist including the relationship and monophyly of different tribes and some genera. The aim of this study was to characterize the mitogenome of Coprophanaeus ensifer in order to establish its position within the Scarabaeidae family and to contribute to the resolution of some phylogenetic uncertainties. The mitogenome was sequenced on the Illumina HiSeq 4000, assembled using the Mitobim software and annotated in MITOS WebServer. The phylogenetic trees were reconstructed by Bayesian inference. The C. ensifer mitogenome is a molecule of 14,964 bp that contains the number and organization of the genes similar to those of most Coleoptera species. Phylogenetic reconstruction suggests monophyly of the tribe Phanaeini and supports the hypothesis that Coprini is a sister group of Phanaeini. The results also revealed the position of the tribe Oniticellini which is grouped with Onthophagini and Onitini. The geographic distribution of these species that form the most ancestral clade suggests with Scarabaeinae originated in Africa. Keywords: Dung beetle, mitochondrial genome, phylogenomics     

Are ball-rolling (Scarabaeini,Gymnopleurini, Sisyphini) and tunnelling scarabaeine dung beetles equally choosy about the size of ingested dung particles?

Abstract.  1. The maximum size of ingested particles was determined in 11 species of ball-rolling, adult dung beetle (Scarabaeidae: Scarabaeinae) by mixing small latex or glass balls of known diameter into their food. The tribes Scarabaeini, Gymnopleurini, and Sisyphini (four, four, and three species respectively) were represented, with mean body sizes ranging from 0.33 to 4.0 g fresh weight.
2. Only particles with maximum diameters of 4–85 µm were ingested. Hence rollers, like other known beetles feeding on fresh dung, filter out larger, indigestible plant fragments and confine ingestion to small particles of higher nutritional value.
3. The maximum diameter of ingested particles increased significantly with body weight, whereas taxon (tribe) had no additional effect. Because big rollers accept larger particles than do tunnellers (which make dung stores for feeding and breeding in the soil immediately below the pat) of similar weight, the slope of the diameter-against-weight regression for rollers was significantly higher than that found earlier for tunnellers.
4. An explanation could be that a typical food ball made by a roller is considerably smaller than the amount of dung available to a feeding tunneller of the same size. If the roller were as choosy about particle size as the tunneller, it might not get enough food. This applies to large rollers in particular because their food balls contain a higher proportion of coarse fibres than those made by small species.     

How localized are the distributions of Australian scarabs (Coleoptera,Scarabaeoidea)?

The degree of localization in the distribution of Australian scarabaeine (320 spp.), aphodiine (160 spp.), geotrupid (166 spp.) and dynastine (182 spp.) scarabs is analysed. Localization differs among these groups, appearing to increase marginally from the Dynastinae, to the Aphodiinae, to the Scarabaeinae, to the Geotrupidae, but with significant differences within groups, e.g. the scarabaeine tribes Onthophagini and Scarabaeini. However, the Australian fauna lacks the high proportion of widespread species characteristic of the West Palaearctic Scarabaeinae, Aphodiinae and Geotrupidae. This difference reflects the different ecological histories of the two regions.     

An old adaptive radiation of forest dung beetles in Madagascar

Adaptive radiations of mammals have contributed to the exceptionally high levels of biodiversity and endemism in Madagascar. Here we examine the evolutionary history of the endemic dung beetle tribe Helictopleurini (Scarabaeidae) and its relationship to the widely distributed Oniticellini and Onthophagini. Helictopleurini species are dependent on mammals for their resources. We date the single origin of the tribe at 37 to 23 MY ago, indicating overseas colonization of Madagascar. The main radiation occurred concurrently with the main radiations of lemurs. The ancestors of Helictopleurini are inferred to have been coprophagous species inhabiting open habitats. Subsequent evolution has involved a shift into forests, changes in resource use to a more generalized diet, and changes in body size. Four species of the extant 65 species have shifted to use the dung of the recently introduced cattle in open habitats, allowing these species to greatly expand their geographical ranges.     

A molecular phylogenetic analysis of the Scarabaeinae (dung beetles)

The dung beetles (Scarabaeinae) include ca. 5000 species and exhibit a diverse array of morphologies and behaviors. This variation presumably reflects the adaptation to a diversity of food types and the different strategies used to avoid competition for vertebrate dung, which is the primary breeding environment for most species. The current classification gives great weight to the major behavioral types, separating the ball rollers and the tunnelers, but existing phylogenetic studies have been based on limited taxonomic or biogeographic sampling and have been contradictory. Here, we present a molecular phylogenetic analysis of 214 species of Scarabaeinae, representing all 12 traditionally recognized tribes and six biogeographical regions, using partial gene sequences from one nuclear (28S) and two mitochondrial (cox1, rrnL) genes. Length variation in 28S (588-621 bp) and rrnL (514-523 bp) was subjected to a thorough evaluation of alternative alignments, gap-coding methods, and tree searches using model-based (Bayesian and likelihood), maximum parsimony, and direct optimization analyses. The small-bodied, non-dung-feeding Sarophorus+Coptorhina were basal in all reconstructions. These were closely related to rolling Odontoloma+Dicranocara, suggesting an early acquisition of rolling behavior. Smaller tribes and most genera were monophyletic, while Canthonini and Dichotomiini each consisted of multiple paraphyletic lineages at hierarchical levels equivalent to the smaller tribes. Plasticity of rolling and tunneling was evidenced by a lack of monophyly (S-H test, p > 0.05) and several reversals within clades. The majority of previously unrecognized clades were geographical, including the well-supported Neotropical Phanaeini+Eucraniini, and a large Australian clade of rollers as well as tunneling Coptodactyla and Demarziella. Only three lineages, Gymnopleurini, Copris+Microcopris and Onthophagus, were widespread and therefore appear to be dispersive at a global scale. A reconstruction of biogeographical characters recovered 38-48 transitions between regions and an African origin for most lineages. Dispersal-vicariance analysis supported an African origin with links to all other regions and little back-migration. Our results provide a new synthesis of global-scale dung beetle evolution, demonstrating the great plasticity of behavioral and morphological traits and the importance of biogeographic distributions as the basis for a new classification.     

Phylogenetic analyses reveal reliable morphological markers to classify mega‐diversity in Onthophagini dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae)

Based largely on homoplastic characters of external morphology, the current systematics of the tribe Onthophagini and allied dung beetle lineages is unstable, contradictory, and thus inefficient. A number of recently proposed molecular phylogenies conflict strongly with each other and with formal classification, and none of them provides new tools for the improvement of dung beetle systematics. We explored the source of these inconsistencies by performing an independent, morphology‐based phylogenetic analysis of the “Serrophorus complex”, one of the most systematically confusing knots among the onthophagines, that involves 52 species from various genera of Onthophagini and allied tribes. The phylogenetic pattern revealed conflicts with existing classifications and with most of the earlier molecular phylogenies. However, it was largely congruent with the molecular phylogeny (Evolution 2005, 59 , 1060), using the largest gene sampling thus far. All current competing phylogenetic hypotheses were evaluated against each other, and the degree of their biogeographic plausibility was used as an additional evaluative criterion. Of the 91 morphological characters involved in our analyses, traits belonging to the endophallic sclerites of the aedeagus had a very strong phylogenetic signal. Terminology of these endophallic characters was established and their morphology was studied in detail, illustrated, and presented as a tool for further practical use. The enormous variety of shapes of the lamella copulatrix within the Onthophagini and allies present a methodological problem in character coding for phylogenetic analyses. Based on the performance of alternative coding approaches, it is argued that a seemingly less informative absence/presence coding scheme would be a better choice. The phylogenetic structure of the Serrophorus complex has been largely resolved, and some taxonomic changes improving its systematics are recommended.
© The Willi Hennig Society 2011.     

Several steps of lateral gene transfer followed by events of ‘birth-and-death’ evolution shaped a fungal sorbicillinoid biosynthetic gene cluster

Dung beetles (subfamily Scarabaeinae) are popular model organisms in ecology and developmental biology, and for the last two decades they have experienced a systematics renaissance with the adoption of modern phylogenetic approaches. Within this period 16 key phylogenies and numerous additional studies with limited scope have been published, but higher-level relationships of this pivotal group of beetles remain contentious and current classifications contain many unnatural groupings. The present study provides a robust phylogenetic framework and a revised classification of dung beetles. We assembled the so far largest molecular dataset for dung beetles using sequences of 8 gene regions and 547 terminals including the outgroup taxa. This dataset was analyzed using Bayesian, maximum likelihood and parsimony approaches. In order to test the sensitivity of results to different analytical treatments, we evaluated alternative partitioning schemes based on secondary structure, domains and codon position. We assessed substitution models adequacy using Bayesian framework and used these results to exclude partitions where substitution models did not adequately depict the processes that generated the data. We show that exclusion of partitions that failed the model adequacy evaluation has a potential to improve phylogenetic inference, but efficient implementation of this approach on large datasets is problematic and awaits development of new computationally advanced software. In the class Insecta it is uncommon for the results of molecular phylogenetic analysis to lead to substantial changes in classification. However, the results presented here are congruent with recent morphological studies and support the largest change in dung beetle systematics for the last 50 years. Here we propose the revision of the concepts for the tribes Deltochilini (Canthonini), Dichotomiini and Coprini; additionally, we redefine the tribe Sisyphini. We provide and illustrate synapomorphies and diagnostic characters supporting the new concepts to facilitate diagnosability of the redefined tribes. As a result of the proposed changes a large number of genera previously assigned to these tribes are now left outside the redefined tribes and are treated as incertae sedis. The present study redefines dung beetles classification and gives new insight into their phylogeny. It has broad implications for the systematics as well as for various ecological and evolutionary analyses in dung beetles.     

Influence of Evolutionary Allometry on Rates of Morphological Evolution and Disparity in strictly Subterranean Moles (Talpinae,Talpidae, Lipotyphla,Mammalia)

The adaptation to a particular function could directly influence the morphological evolution of an anatomical structure as well as its rates. The humeral morphology of moles (subfamily Talpinae) is highly modified in response to intense burrowing and fully fossorial lifestyle. However, little is known of the evolutionary pathways that marked its diversification in the two highly fossorial moles tribes Talpini and Scalopini. We used two-dimensional landmark-based geometric morphometrics and comparative methods to understand which factors influenced the rates and patterns of the morphological evolution of the humerus in 53 extant and extinct species of the Talpini (22 extant plus 12 extinct) and Scalopini (six extant plus 13 extinct) tribes, for a total of 623 humeri. We first built a synthetic phylogeny of extinct and extant taxa of the subfamily Talpinae based on all the available information from known phylogenies, molecular data, and age ranges of fossil records. We tested for evolutionary allometry by means of multivariate regression of shape on size variables. Evolutionary allometric trajectories exhibited convergence of humeral shape between the two tribes, even when controlling for phylogeny, though a significant differences in the evolutionary rates was found between the two tribes. Talpini, unlike Scalopini, seem to have reached a robust fossorial morphology early during their evolution, and their shape disparity did not change, if it did not decrease, through time. Furthermore, the basal Geotrypus spp. clearly set apart from the other highly fossorial moles, exhibiting a significant acceleration of evolutionary shifts toward higher degree of fossorial adaptation. Our observations support the hypothesis that the evolution of allometry may reflect a biological demand (in this case functional) that constrains the rates of evolution of anatomical structures.     

Mitochondrial phylogenomics reveals insights into taxonomy and evolution of Penaeoidea (Crustacea: Decapoda)

The taxonomy and phylogeny of Penaeoidea have long been fraught with controversy. Here, we carried out the first mitochondrial phylogenomic analysis on all the penaeoid families and tribes, including nine newly sequenced and 14 published mitogenomes, towards elucidating the phylogeny and evolutionary history of Penaeoidea. All these nine mitogenomes exhibit the pancrustacean ground pattern, except that Benthonectes filipes contains two additional clusters of tRNA^Ala, tRNA^Arg and tRNA^Asn and an uncommon noncoding region. The resulted phylogenetic tree is generally well resolved with Benthesicymidae sister to Aristeidae, forming a clade with Solenoceridae. Contrary to traditional classification, this clade has a sister relationship with the tribe Penaeini of the family Penaeidae. The family Sicyoniidae is deeply nested within the penaeid tribe Trachypenaeini which forms a sister clade with the remaining penaeid tribe, Parapenaeini. As the family Penaeidae is recovered to be polyphyletic, the three tribes in Penaeidae are all elevated to familial status. On the other hand, the family Sicyoniidae is retained to accommodate Trachypenaeini because they are now synonyms and the former name is more senior. This work is the first molecular analysis concurring with the latest findings in fossil assessments showing that Parapeaneini is the most primitive in Penaeoidae. Our results also illustrate a shallow‐water origin and an onshore–offshore evolutionary shift in penaeoid shrimps.     

Primary classification and phylogeny of the Polemoniaceae, with comments on molecular cladistics

The system of classification of the Polemoniaceae currently in use was published by Grant in 1959. Much new evidence concerning relationships in the family has been obtained by numerous workers since 1959, and the old system is in need of revision. A revised system down to the genus level, based on conventional and unconventional characters, including molecular evidence, is presented here. Nineteen genera are grouped into eight tribes and two subfamilies. Three new tribes are described: Acanthogilieae, Loeselieae, and Leptodactyloneae. Several genera are transferred to new groups. The phylogeny of the family is discussed in the light of both the older and new evidence. The approach used in constructing both the 1959 and new systems is that of evolutionary systematics. Two recent (1996, 1997) family-wide surveys of cpDNA and rDNA use cladistic methods of analysis to arrive at sets of major groups. Some of this molecular evidence has been adopted for the present revised system. However, much incongruence still exists between the new sets of clades, on the one hand, and the present revised system or the still-viable parts of the 1959 system on the other hand. The incongruences call for an examination and comparison of the contrasting methods of evolutionary systematics and molecular cladistics. A fundamental flaw in the 1996 and 1997 treatments is the attempt to classify plants on the basis of single-gene gene trees.     

Molecular and morphological phylogeny of Menispermaceae (Ranunculales)

The phylogeny of Menispermaceae (Ranunculales) is reconstructed here using both morphological and molecular data from a broad sample of species. Morphological data include characters of leaves, wood, flowers, fruits, seeds, pollen and phytochemistry from 73 species representing the different subgroups recognized within the family. This dataset allowed us to study the morphological evolutionary trends in Menispermaceae. The molecular data focused on cpDNA sequences: rbcL and atpB. Maximum parsimony (MP) and Bayesian methods were used to reconstruct the phylogeny of Menispermaceae. The results obtained from the three datasets are partly incongruent. The morphology indicates that Menispermaceae can be divided into two major groups, one including the tribes Fibraureae and Tinosporeae, the other one including the tribes Anomospermeae, Pachygoneae and Menispermeae. Only Burasaia Thouars was not included in any of these groups. The characters of fruits and seeds appeared to be most useful to differentiate these groups whereas convergences were found for the androecium.     

Molecular phylogeny and evolutionary habitat transition of the flower bugs (Heteroptera: Anthocoridae)

We performed a molecular phylogenetic study of the Anthocoridae, the flower bugs, based on maximum likelihood, maximum parsimony, and Bayesian analyses of ～ 3000 base pairs (bp) of DNA sequence from the mitochondrial 16S rRNA and nuclear 18S rRNA and 28S rRNA genes for 44 taxa. Our phylogenetic analyses indicates that (i) the tribe Cardiastethini (Dufouriellini) could be a paraphyletic group, as the genera Amphiareus and Dysepicritus are not included in the tribe; (ii) the main subgroups, Oriini and Anthocorini, are monophyletic within Anthocoridae; (iii) three tribes of Blaptostethini, Xylocorini, and Scolopini are separated from the main anthocorid clade which is composed of Anthocorini, Cardiastethini, and Oriini, suggesting that Anthocoridae could not be monophyletic. We compared our molecular phylogeny to previous hypotheses of evolutionary relationships within Cimicoidea based on different anthocorid classification systems using alternative hypothesis tests (Kishino-Hasegawa and Shimodaira-Hasegawa tests). BayesTraits were used to examine the ancestral character states inferring historical habitat patterns of the Anthocoridae. Reconstruction of the ancestral habitat patterns of the Anthocoridae suggests that dead plants may have served as an important habitat for the common ancestor of anthocorids. The biological events such as diversification of angiosperms and anthocorid prey might have provided anthocorids with more habitat options, such as living plants; thereafter, Anthocorini and Amphiareus appeared to have evolved increasingly specialized habitat relationships.     

A molecular phylogeny of the Cephinae (Hymenoptera, Cephidae) based on mtDNA COI gene: a test of traditional classification

Cephinae is traditionally divided into three tribes and about 24 genera based on morphology and host utilization. There has been no study testing the monophyly of taxa under a strict phylogenetic criterion. A molecular phylogeny of Cephinae based on a total of 68 sequences of mtDNA COI gene, representing seven genera of Cephinae, is reconstructed to test the traditional limits and relationships of taxa. Monophyly of the traditional tribes is not supported. Monophyly of the genera are largely supported except for Pachycephus. A few host shift events are suggested based on phylogenetic relationships among taxa. These results indicate that a more robust phylogeny is required for a more plausible conclusion. We also report two species of Cephus for the first time from Turkey.     

Evolutionary strategies of human T-cell lymphotropic virus type II

Human T-cell lymphotropic virus type II (HTLV-II) primarily infects two different populations in which the virus is transmitted in very diverse ways. In endemically infected populations, the virus is propagated through sexual contact, and by mother to child transmission via breast-feeding, among intravenous drug users (IDUs), spread is mainly due to blood-borne transmission via needle sharing. The phylogeny of HTLV-II strains isolated from American Indian and Pygmy tribes and strains from IDUs, reveal that the virus originated on the African continent as a result of a simian to human transmission at least 400,000 years ago. HTLV-II was very likely introduced into the American continent during one or more migrations of HTLV-II infected Asian populations over the Bering land bridge, some 15,000-35,000 years ago. During the last few decades, HTLV-II has been transmitted from native American Indians to IDUs at least twice, followed by a rapid spread of the virus in the drug users population world-wide due to the practice of needle sharing. Molecular clock analysis showed that HTLV-II has two different evolutionary rates, with the molecular clock for the virus in IDUs ticking 150-350 times faster than the one in endemically infected tribes: 2.7x10(-4) compared to 1.7/7.3x10(-7) nucleotide substitutions per site per year in the LTR region. Although many of the HTLV-II infected drug users are co-infected with HIV, the dramatic acceleration of the evolutionary rate seems to be mainly related to the different modes of transmission in the two populations. These contrasting evolutionary rates correlate with an endemic spread of HTLV-II in infected tribes compared to an epidemic spread in IDUs.     

Oil-producing flowers within the Iridoideae (Iridaceae): evolutionary trends in the flowers of the New World genera

Background and Aims

Oil-producing flowers related to oil-bee pollination are a major innovation in Neotropical and Mexican Iridaceae. In this study, phylogenetic relationships were investigated among a wide array of New World genera of the tribes Sisyrinchieae, Trimezieae and Tigridieae (Iridaceae: Iridoideae) and the evolution of floral glandular structures, which are predominantly trichomal elaiophores, was examined in relation to the diversification of New World Iridaceae.

Methods

Phylogenetic analyses based on seven molecular markers obtained from 97 species were conducted to produce the first extensive phylogeny of the New World tribes of subfamily Iridoideae. The resulting phylogenetic hypothesis was used to trace the evolutionary history of glandular structures present in the flowers of numerous species in each tribe. Hypotheses of differential diversification rates among lineages were also investigated using both topological and Binary-State Speciation and Extinction methods.

Key Results and Conclusions

Floral glandular structures and especially trichomal elaiophores evolved multiple times independently in the American tribes of Iridoideae. The distribution pattern of species displaying glandular trichomes across the phylogeny reveals lability in the pollination system and suggests that these structures may have played a significant role in the diversification of the Iridoideae on the American continent.     

Systematics of the dung beetle tribe Sisyphini Mulsant (Scarabaeidae: Scarabaeinae) inferred from a molecular phylogeny and biogeography of southern African species

The tribe Sisyphini Mulsant was recently redefined following the transfer of the endemic southern African genus Epirinus Dejean from the polyphyletic tribe Deltochilini Lacordaire. A molecular phylogeny of the southern African members of Sisyphini supports Epirinus as sister to Sisyphus Latreille and recovered three major clades in Sisyphus classified here as subgenera Sisyphus (Neosisyphus Müller) stat. rev. , Sisyphus (Parasisyphus Barbero, Palestrini & Zunino) stat.n. and Sisyphus (Sisyphus) stat.n. A molecular clock analysis suggests that Sisyphus and Epirinus diverged from their last common ancestor during the Lower to Middle Oligocene (c. 29.37 Ma). Biogeographical analysis indicated that southern African Sisyphus species are centred in the east and northeast in Highveld grassland and warmer savannah regions. By contrast, Epirinus species are largely restricted to the southwest and southeast in the cooler winter and bimodal rainfall regions plus arid highland Karoo and Highveld grasslands. Based on morphological and biogeographical differences between Epirinus and Sisyphus, we propose that the monogeneric Epirinus be placed in its own tribe, Epirinini van Lansberge stat. rev.     

Molecular phylogeny of the Chinese ranids inferred from nuclear and mitochondrial DNA sequences

The phylogeny of representative species of Chinese ranids was reconstructed using two nuclear (tyrosinase and rhodopsin) and two mitochondrial (12S rRNA, 16S rRNA) DNA fragments. Maximum parsimony, Bayesian, and maximum likelihood analyses were employed. In comparison with the results from nuclear and mitochondrial data, we used nuclear gene data as our preferred phylogenetic hypothesis. We proposed two families (Ranidae, Dicroglossidae) for Chinese ranids, with the exception of genus Ingerana. Within Dicroglossidae, four tribes were supported including Dicroglossini, Paini, Limnonectini, and Occidozygini. A broader sampling strategy and evidence from additional molecular markers are required to decisively evaluate the evolutionary history of Chinese ranids.