Integrative taxonomy reveals extreme morphological conservatism in sympatric Mugil species from the Tropical Southwestern Atlantic

The biodiversity crisis has had particularly harsh impacts on marine environments. However, there is still considerable uncertainty about how many species have been seriously impacted and the effectiveness of protection measures (e.g., marine protected areas or MPAs) due to high levels of cryptic species in many taxa. Here, we employ an integrative taxonomy approach to mullet species in the genus Mugil. In addition to its high economic value, this genus is notable for having diversified ~29 million years ago without marked morphological and ecological divergence. We obtained 129 specimens of Mugil from the Coral Coast MPA, the largest of its kind in the Tropical Southwestern Atlantic marine province. Although morphometric and meristic traits revealed six taxonomically recognized species, only five mitochondrial lineages were observed. All individuals morphologically identified as M. incilis belonged to the mitochondrial lineage of Mugil curema, which is consistent with misidentification of morphologically similar species and an overestimation of species diversity. Remarkably, Mugil species in our sample that diverged up to ~23 million years ago are also the most morphologically similar (M. curema and M. rubrioculus), suggesting extreme morphological conservatism, possibly driven by similarities in habitat use and life‐history traits. This study demonstrates the potential utility of integrative taxonomy (including DNA barcoding) for contributing to the conservation and sustainable use of natural resources.     

Molecular phylogeny endorses the relationship between carnivorous and filter‐feeding tunicates (Tunicata,Ascidiacea)

Tatián, M., Lagger, C., Demarchi, M. & Mattoni, C. (2011). Molecular phylogeny endorses the relationship between carnivorous and filter‐feeding tunicates (Tunicata, Ascidiacea).—Zoologica Scripta, 40, 603–612. The phylogeny of the tunicates (animals considered the closest relatives to the vertebrates) is not yet completely defined, especially the evolutionary relationships within the class. Molecular studies do not include particular benthic deep‐sea species that show morphological changes in the evolution from filter feeding into a carnivorous‐feeding habit. According only to morphological features, these animals are considered as a part of the Class Ascidiacea (Family Hexacrobylidae), but also as a different class, Sorberacea, belonging to the Phylum Tunicata. In this study, we present a phylogenetic analysis based on 18S rDNA sequences, which clearly included these animals in Ascidiacea but in the Family Molgulidae, faster‐evolving ascidians with a high evolution rate. This finding supports the idea that carnivory in Molgulidae represents a more recent adaptation to life in the ocean deep bottoms, where organisms have to adapt themselves to a less plentiful particulate organic carbon supply. Based on molecular and morphological evidence, we propose the following new synonymy: Hexacrobylidae Seeliger 1906 = Molgulidae Lacaze‐Duthiers, 1877 .     

MORPHOLOGICAL EVOLUTION OF THE DROSOPHILA VIRILIS SPECIES GROUP AS ASSESSED BY RATE TESTS FOR NATURAL SELECTION ON QUANTITATIVE CHARACTERS

Two rate tests for assessing natural selection on quantitative traits are discussed for their usefulness in macroevolutionary and adaptational studies. The underlying assumptions and parameter estimation for the constant-heritability (CH) and mutation-drift-equilibrium (MDE) models, which are the bases for these tests, are discussed. The purpose of these rate tests is to determine whether morphological change has occurred either too fast to be explained by neutral drift, which suggests directional selection, or too slow, which suggests stabilizing selection. Previous formulations of these rate tests have ignored the phylogenetic component. Several models of evolution are considered to help account for phylogeny in the context of rate tests. The MDE rate test for stabilizing selection was performed on nine morphological characters among several species of the Drosophila virilis species group. These tests can be interpreted to suggest that stabilizing selection has probably been a major factor in producing the observed similarity among the Drosophila species examined.     

Chromosome and breeding system evolution of the genus Mercurialis (Euphorbiaceae): implications of ITS molecular phylogeny

 The internal transcribed spacers (ITS1 and ITS2) of nuclear ribosomal DNA were amplified and sequenced from 19 samples representing all species of the genus Mercurialis and two outgroup species, Ricinus communis and Acalypha hispida. The length of ITS1 in the ingroups ranged from 223 to 246 bp and ITS2 from 210 to 218 bp. Sequence divergence between pairs of species ranged from 1.15% to 25.88% among the ingroup species in the combined data of ITS1 and ITS2. Heuristic phylogenetic analyses using Fitch parsimony on the combined data of ITS1 and ITS2 with gaps treated as missing generated 45 equally parsimonious trees. The strict consensus tree was principally concordant with morphological classification. Within the genus, the ITS sequences recognised two main infrageneric clades: the M. perennis complex including three Eurasian stoloniferous species (M.␣leiocarpa, M. ovata and M. perennis) and the western Mediterranean group including eight both annual and perennial species. Of the western Mediterranean clade, the annual and perennial species grouped respectively into two different groups, and the annual life form is revealed as a synapomorphic character derived from perennial, whereas in the Eurasian clade ITS phylogeny suggested M. leiocarpa as basal clade sister to M.␣perennis and M. ovata. ITS phylogeny failed to resolve the relationships among the different cytotypes of M. ovata and M. perennis. ITS phylogeny also suggested rapid karyotypic evolution for the genus. The karyotypic divergence among the perennial species of western Mediterranean region did not corroborate the nucleotide sequence divergence among the species. Optimisation of chromosome numbers onto the ITS phylogeny suggested x=8 to be the ancestral basic chromosome number of the genus. ITS phylogeny confirmed that the androdioecy of M. ambigua is derived from dioecy. The nucleotide heterozygosity and additivity in ITS sequences clearly confirm the interspecific hybridisation in the genus Mercurialis. Received December 22, 2001; accepted May 21, 2002?Published online: November 14, 2002 Address of the authors: Martin Kr?henbühl, Yong-Ming Yuan (correspondence) and Philippe Küpfer, Institut de Botanique, Laboratoire de botanique évolutive, Université de Neuchatel, Emile-Argand 11, CH-2007 Neuchatel, Suisse. (e-mail: yong-ming.yuan@unine.ch)     

Molecular phylogeny of Mugilidae fishes revised

Systematics derived from morphological characters often does not correspond with the evolutionary processes underlying the divergence within a group of organisms. In the family Mugilidae (Teleostei) morphological similarities have resulted in inconsistencies between taxonomy and phylogeny among its species, and particularly for the genera Mugil, Liza and Chelon where both intrageneric and intergeneric phylogenetic clarifications are needed. To address these issues, the direct sequencing of the mitochondrial region that encodes Phenylalanine (69 bp), 12S rRNA (842 bp), cytochrome c oxidase subunit I (651 bp) and cytochrome b (702 bp) was carried out. The data reveal that Mugil platanus and Mugil liza represent a continuum of a single species, closely related to but distinct from Mugil cephalus which itself appears to comprise a grouping of multiple and closely related species. This species complex was genetically distinct from Mugil curema, which, based on three clearly diverged species identified in this study along the Atlantic coast of the Americas, requires extensive taxonomic revision throughout its world-wide distribution. Unlike the monophyly supported within Mugil, relationships within Liza are paraphyletic, and a taxonomic revision of the genera Liza, Chelon and Oedalechilus is needed.     

Patterns of evolution in western North American Mimulus (Phrymaceae)

A well-supported phylogeny is presented from both chloroplast DNA (the trnL/F region) and two regions of nuclear rDNA (ITS [internal transcribed spacer] and ETS [external transcribed spacer]) with nearly complete sampling for Mimulus (Phrymaceae) in western North America. Three separate genera are derived from within the clade that contains all the Mimulus species in western North America. The taxonomic status of the proposed sections of Mimulus and the relationships of many taxonomically difficult species are considered with observations on morphological evolution. Discordance between data sources provides support for the hypothesis that M. evanescens is a hybrid between M. latidens and M. breviflorus. In two major clades (Eunanus and Diplacus), patterns of genetic variation do not match the current taxonomy. The clustering of taxa in Eunanus is strongly associated with geographic distributions. Mimulus aurantiacus sensu Thompson, M. nanus, and M. floribundus are found to be progenitor species to other species that appear to be derived from within them. Polyploidy and aneuploidy events are clustered near the tips of the phylogeny. Thus, these two mechanisms are concluded to have played a relatively small role in the evolution of persistent lineages in Mimulus. The phylogenetic distribution of rare taxa is also examined.     

Phylogenies from genetic and morphological characters do not support a revision of Gigasporaceae (Glomeromycota) into four families and five genera

The family Gigasporaceae consisted of the two genera Gigaspora and Scutellospora when first erected. In a recent revision of this classification, Scutellospora was divided into three families and four genera based on two main lines of evidence: (1) phylogenetic patterns of coevolving small and large rRNA genes and (2) morphology of spore germination shields. The rRNA trees were assumed to accurately reflect species evolution, and shield characters were selected because they correlated with gene trees. These characters then were used selectively to support gene trees and validate the classification. To test this new classification, a phylogenetic tree was reconstructed from concatenated 25S rRNA and β-tubulin gene sequences using 35% of known species in Gigasporaceae. A tree also was reconstructed from 23 morphological characters represented in 71% of known species. Results from both datasets showed that the revised classification was untenable. The classification also failed to accurately represent sister group relationships amongst higher taxa. Only two clades were fully resolved and congruent among datasets: Gigaspora and Racocetra (a clade consisting of species with spores having one inner germinal wall). Other clades were unresolved, which was attributed in part to undersampling of species. Topology of the morphology-based phylogeny was incongruent with gene evolution. Five shield characters were reduced to three, of which two were phylogenetically uninformative because they were homoplastic. Therefore, most taxa erected in the new classification are rejected. The classification is revised to restore the family Gigasporaceae, within which are the three genera Gigaspora, Racocetra, and Scutellospora. This classification does not reflect strict topology of either gene or morphological evolution. Further revisions must await sampling of additional characters and taxa to better ascertain congruence between datasets and infer a more accurate phylogeny of this important group of fungi.     

Molecular phylogeny and evolution of phenotype in silica‐scaled chrysophyte genus Mallomonas

The evolution of phenotypes is highly understudied in protists, due to the dearth of morphological characters, missing fossil record, and/or unresolved phylogeny in the majority of taxa. The chrysophyte genus Mallomonas (Stramenopiles) forms species‐specific silica scales with extraordinary diversity of their ornamentation. In this paper, we molecularly characterized three additional species to provide an updated phylogeny of 43 species, and combined this with evaluations of 24 morphological traits. Using phylogenetic comparative methods, we evaluated phylogenetic signal in traits, reconstructed the trait evolution, and compared the overall phylogenetic and morphological diversity. The majority of traits showed strong phylogenetic signal and mostly dynamic evolution. Phylogenetic relatedness was often reflected by the phenotypic similarity. Both V‐rib and dome are very conservative structures that are presumably involved in precise scale overlap and bristle attachment, respectively. Based on modern species, it seems the dome firstly appeared on apical and/or caudal scales, and only later emerged on body scales. Bristle was presumably present in the common ancestor and gradually elongated ever since. However, most other morphological traits readily changed during the evolution of Mallomonas.     

Exploring the evolution of humus collecting leaves in drynarioid ferns (Polypodiaceae, Polypodiidae) based on phylogenetic evidence

Most species of the paleotropic fern genera Aglaomorpha and Drynaria, together constituting a monophyletic clade (drynarioid ferns), possess humus-collecting structures as an adaptation to their epiphytic life form. Humus-collectors are either present as a specialized foliar structure (external leaf dimorphism) or as a specialized leaf part (internal dimorphism). Apart from these basic patterns there are several forms of reduction and an internal fertile – sterile dimorphism in Aglaomorpha. We present a phylogeny of drynarioid ferns based on morphological and molecular (cpDNA) markers. The genus Aglaomorpha was found to be monophyletic, whereas Drynaria is likely to be a paraphyletic assemblage including a grade of Himalayan to Southern Chinese taxa basal to Aglaomorpha. The evolution of humus-collectors is reconstructed by plotting their character state changes onto the obtained phylogeny. Despite the complex morphological pattern across species, evolution of drynarioid humus-collecting structures can be reconstructed postulating a simple sequence of character state changes based on only a few elementary processes.     

Taxonomic Study of Korean <Emphasis Type="Italic">Scirpus</Emphasis> L. s.l. (Cyperaceae) II: Pattern of Phenotypic Evolution Inferred from Molecular Phylogeny

Scirpus L. s.l. is well known as one of the polyphyletic groups in Cyperaceae. Recent molecular phylogenetic studies clearly suggested that Scirpus s.l. should be separated into several independent genera. In this study, we intend to present the morphological variations and patterns of phenotypic evolution based on molecular phylogeny of Korean Scirpus s.l. Five genera, including 21 taxa from Korean Scirpus s.l., were examined: three species of Bolboschoenus, three species of Schoenoplectus, eight species of Schoenoplectiella including one hybrid, five species of Scirpus, and two species of Trichophorum. For morphological analyses, 23 and 48 characters were selected from vegetative and reproductive organs, respectively. Molecular phylogeny was inferred from a nuclear ribosomal internal transcribed spacer, the chloroplast rbcL gene, and the chloroplast trnL-F region. Distinct characteristics and quantitative variation was presented for identification of the five genera and their species. A pronounced pattern of morphological character change was reduction, although many other character states seem to be homoplastic. We suggest that the reduction of phenotypic characteristics has been expressed in terms of condensation of internodes, reduction of leaf blade, and simplification of inflorescence among five genera of Korean Scirpus s.l.     

Molecular evolution and phylogeny of theDrosophila virilis species group as inferred by two-dimensional electrophoresis

Summary Systematic relationship among the 12 species of theDrosophila virilis species group, andDrosophila robusta, were investigated by the use of two-dimensional electrophoresis (2-DE). A total of 389 protein characters (about 200 loci) were scored and analyzed both phylogenetically and phenetically. The resulting phylogeny was found to be largely concordant with the current views of evolution among these species based on other independent morphological, chromosomal, electrophoretic, and immunological data sets, although some notable differences were observed. The 2-DE data also appeared to be useful for constructing a molecular clock to date the absolute times of divergence among the species. It appears from this analysis that the evolution of the major clades within the species group occurred about 20 million years ago. Previous suggestions that the rate of molecular evolution was different between the virilis and montana phylads was not confirmed. The technique of 2-DE seems to be an excellent tool for reconstructing phylogenies and should be particularly valuable for examining relatively closely related species.     

Molecular phylogeny of hinge‐beak shrimps (Decapoda: Caridea: Rhynchocinetes and Cinetorhynchus) and allies: a formal test of familiar and generic monophyly using a multilocus phylogeny

The Rhynchocinetidae (‘hinge‐beak’ shrimps) is a family of marine caridean decapods with considerable variation in sexual dimorphism, male weaponry, mating tactics, and sexual systems. Thus, this group is an excellent model with which to analyse the evolution of these important characteristics, which are of interest not only in shrimps specifically but also in animal taxa in general. Yet, there exists no phylogenetic hypothesis, either molecular or morphological, for this taxon against which to test either the evolution of behavioural traits within the Rhynchocinetidae or its genealogical relationships with other caridean taxa. In this study, we tested (1) hypotheses on the phylogenetic relationships of rhynchocinetid shrimps, and (2) the efficacy of different (one‐, two‐, and three‐phase) methods to generate a reliable phylogeny. Total genomic DNA was extracted from tissue samples taken from 17 species of Rhynchocinetidae and five other species currently or previously assigned to the same superfamily (Nematocarcinoidea); six species from other superfamilies were used as outgroups. Sequences from two nuclear genes (H3 and Enolase) and one mitochondrial gene (12S) were used to construct phylogenies. One‐phase phylogenetic analyses (SATé‐II) and classical two‐ and three‐phase phylogenetic analyses were employed, using both maximum likelihood and Bayesian inference methods. Both a two‐gene data set (H3 and Enolase) and a three‐gene data set (H3, Enolase, 12S) were utilized to explore the relationships amongst the targeted species. These analyses showed that the superfamily Nematocarcinoidea, as currently accepted, is polyphyletic. Furthermore, the two major clades recognized by the SATé‐II analysis are clearly concordant with the genera Rhynchocinetes and Cinetorhynchus, which are currently recognized in the morphological‐based classification (implicit phylogeny) as composing the family Rhynchocinetidae. The SATé‐II method is considered superior to the other phylogenetic analyses employed, which failed to recognize these two major clades. Studies using more genes and a more complete species data set are needed to test yet unresolved inter‐ and intrafamilial systematic and evolutionary questions about this remarkable clade of caridean shrimps. © 2014 The Linnean Society of London     

Nuclear and mitochondrial phylogenies provide evidence for four species of Eurasian badgers (Carnivora)

Del Cerro, I., Marmi, J., Ferrando, A., Chashchin, P., Taberlet, P. & Bosch, M. (2010). Nuclear and mitochondrial phylogenies provide evidence for four species of Eurasian badgers (Carnivora). —Zoologica Scripta, 39, 415–425. The Eurasian badgers (Meles spp.) have a fairly widespread distribution in the Palearctic region and their great morphological variability throughout the vast geographic area has nourished an intense debate about the classification of this taxon. Therefore, the aim of this study was to clarify controversies in Eurasian badger taxonomy by means of a new molecular phylogeny. One‐hundred and seventeen individuals of Eurasian badgers from 18 countries throughout Eurasia were sequenced for up to 3257 bp of nuclear DNA over six loci (ACTC, BGN, CFTR, CHRNA1, TS and TTR) and 512 bp of the mitochondrial DNA control region. Statistical and phylogenetic analyses for combined nDNA, mtDNA and the total‐evidence data clearly showed a strong genetic differentiation in four well‐supported clades, three of which corresponded to allopatric badger species previously defined according to morphological data: Meles meles Linnaeus, 1758 in Europe; Meles leucurus Hodgson, 1847 in the continental part of Asia, except the south‐west part; and M. anakuma Temminck, 1844 in Japan. Up to now, the fourth clade, made up of individuals from south‐west Asia, had been considered as a subspecies. Supported by several pieces of morphological evidence, the new phylogeny revealed that it is necessary to revise the current taxonomic classification of Meles spp. and suggested that the badgers from south‐west Asia should be recognised as a separate species, being renamed M. canescens Blanford, 1875.     

Phylogeny, diversity, and distribution inExostema (Rubiaceae): Implications of morphological and molecular analyses

The neotropical genusExostema comprises 25 species of trees and shrubs, ranging in distribution from Bolivia to Mexico and throughout the West Indies, with most species endemic to the Greater Antilles. Infrageneric relationships and species-level patterns of evolution were investigated in phylogenetic analyses using morphological, molecular, and combined data sets. All data sets resolved three main species groups which correspond to the three sections recognized byMcDowell (1996). However, the analyses of ITS sequence data placed the two South American species basal to the three main clades. Otherwise, the morphological and molecular data are highly compatible, and produce a more robust yet consistent phylogeny in the combined data analysis. Morphological evolution inExostema involves many specializations for xeric habitats, reflecting repeated ecological shifts from moist forest to exposed, seasonally dry environments during the diversification of the genus. Both moth and bee pollination syndromes are found inExostema, and shifts in pollination ecology appear pivotal to the differentiation of the three sections. Biogeographically,Exostema likely originated in South America and migrated via Central America to the Greater Antilles, where the morphological diversification and speciation are most extensive.     

Towards a phylogenetic framework for the evolution of shakes, rattles, and rolls in Myiarchus tyrant-flycatchers (Aves: Passeriformes: Tyrannidae)

A phylogeny of 19 of the 22 currently recognized species of Myiarchus tyrant-flycatchers is presented. It is based on 842bp of mitochondrial DNA (mtDNA) sequences from the ATPase subunit 8 and ATPase subunit 6 genes. Except for the morphologically distinct M. semirufus, mtDNAs of the remaining 18 species fall into either of two clades. One comprises predominantly Caribbean and Central and North American taxa (Clade I), and the other is of predominantly South American taxa (Clade II). The phylogeny is only very broadly concordant with some vocal characters and also with the limited morphological diversity for which the group is well known. Paraphyly in several species (M. swainsoni, M. tuberculifer, M. ferox, M. phaeocephalus, M. sagrae, M. stolidus) suggests that morphological evolution, albeit resulting in limited morphological diversity, has been more rapid than that of mtDNA, or that current taxonomy is faulty, or both. A South American origin for Myiarchus is likely. Dispersal and vicariance both appear to have been involved in generating the present-day distribution of some species. Relatively recent dispersal events out of South America are inferred to have brought species of Clades I and II into broad sympatry. Jamaica has been colonized independently at least twice by members of Clades I and II. The phylogeny brings a historical perspective that in turn suggests that ecological study of closely related species from within each major clade where they are sympatric will be especially rewarding.     

Phylogeny of the Heterodoxus octoseriatus group (Phthiraptera: Boopiidae) from rock-wallabies (Marsupialia: Petrogale)

The phylogenetic position of the Heterodoxus octoseriatus group is inferred from morphological characters. Two character states support monophyly of this group. Another indicates that its sister-group is a group of 11 other Heterodoxus species (here called the H. calabyi group), that infest at least seven other genera of macropodid marsupials. Fourteen potential apomorphies, associated with the male and female genitalia, are identified. Evident rapid and divergent evolution of the genitalia, however, precludes determination of their polarity by comparison with an out-group (the sister-group). Consequently, phylogeny in the H. octoseriatus group is inferred from the close similarity of morphological characters. In light of the phylogenetic analysis and a phenogram, speciation and the evolution of morphological characters in the H. octoseriatus group is discussed.     

A molecular phylogeny for digger wasps in the tribe Ammophilini (Hymenoptera,Apoidea, Sphecidae)

The evolution of parental care strategies in aculeate (stinging) wasps and bees has been much studied from a functional perspective, but relatively little phylogenetic information is available to place this in a rigorous historical context, especially at the species level. We used mitochondrial cytochrome oxidase I and two nuclear genes, the elongation factor‐1α and LW rhodopsin, to investigate the phylogeny of Sphecidae digger wasps. We focus particularly on the tribe Ammophilini, a clade of nonsocial apoid wasps that exhibit unusually diverse parental care strategies. We analysed a 2232 bp dataset for 40 ammophilines plus nine other taxa from within the remaining Sphecidae. Our Bayesian phylogeny provides strong support for the monophyly of Ammophilini and for the monophyly of all six individual ammophiline genera, except that the position of P. affinis within the genus Podalonia is only weakly supported. The monophyly of some, but not all, previously designated species groups within the genus Ammophila is supported. We discuss the implications of our results for the evolution of morphological traits used previously in ammophiline systematics.     

Ecological and morphological attributes of parthenogenetic Japanese Schwiebea species (Acari: Acaridae)

We examined life history traits and spermathecal morphology of both sexual and thelytokous Schwiebea mite species to determine ecological and morphological attributes during the evolution of parthenogenesis in this lineage. We reconstructed a molecular phylogeny of eight Japanese species using the internal transcribed spacer 1 (ITS1) of the ribosomal DNA (rDNA) and compared the sex ratio, developmental period, and egg number (fecundity) of each species within a species group by rearing them in the laboratory. Habitat preference was also analyzed from both collection and literature data. The reconstructed molecular phylogeny suggested that parthenogenesis evolved independently multiple times in this lineage. There were three clusters in the tree, in each of which the idiosoma, leg, setae, and spermathecal morphology of females was similar or identical; this suggested that mites in the same cluster were sister species. There was no relationship between sexual mode and life history traits or habitat preference. These results suggest that sexual and asexual species use different microhabitats. Because S. similis (sexual), S. elongata (thelytokous), and S. estradai (thelytokous) were in the same cluster and spermathecae of the first two were similar while that of the last was distinctively reduced, we hypothesized that speciation occurred in this order and that spermathecae are reduced and eventually lost during the course of parthenogenetic evolution.     

ACCELERATED EVOLUTION OF LAND SNAILS MANDARINA IN THE OCEANIC BONIN ISLANDS: EVIDENCE FROM MITOCHONDRIAL DNA SEQUENCES

An endemic land snail genus Mandarina of the oceanic Bonin (Ogasawara) Islands shows exceptionally rapid evolution not only of morphological and ecological traits, but of DNA sequence. A phylogenetic relationship based on mitochondrial DNA (mtDNA) sequences suggests that morphological differences equivalent to the differences between families were produced between Mandarina and its ancestor during the Pleistocene. The inferred phylogeny shows that species with similar morphologies and life habitats appeared repeatedly and independently in different lineages and islands at different times. Sequential adaptive radiations occurred in different islands of the Bonin Islands and species occupying arboreal, semiarboreal, and terrestrial habitat arose independently in each island. Because of a close relationship between shell morphology and life habitat, independent evolution of the same life habitat in different islands created species possesing the same shell morphology in different islands and lineages. This rapid evolution produced some incongruences between phylogenetic relationship and species taxonomy. Levels of sequence divergence of mtDNA among the species of Mandarina is extremely high. The maximum level of sequence divergence at 16S and 12S ribosomal RNA sequence within Mandarina are 18.7% and 17.7%, respectively, and this suggests that evolution of mtDNA of Mandarina is extremely rapid, more than 20 times faster than the standard rate in other animals. The present examination reveals that evolution of morphological and ecological traits occurs at extremely high rates in the time of adaptive radiation, especially in fragmented environments.