The phylogenetic basis of sexual size dimorphism in orb-weaving spiders (Araneae,Orbiculariae)

Extreme sexual body size dimorphism (SSD), in which males are only a small fraction of the size of the females, occurs only in a few, mostly marine, taxonomic groups. Spiders are the only terrestrial group in which small males are relatively common, particularly among orb-weavers (especially in the families Tetragnathidae and Araneidae) and crab spiders (Thomisidae). We used a taxonomic sample of 80 genera to study the phylogenetic patterns (origins and reversals) of SSD in orb-weaving spiders (Orbiculariae). We collected and compiled male and female size data (adult body length) for 536 species. Size data were treated as a continuous character, and ancestral sizes, for males and females separately, were reconstructed by using Wagner parsimony on a cladogram for the 80 genera used in this study. Of these 80 genera, 24 were female-biased dimorphic (twice or more the body length of the male); the remaining 56 genera were monomorphic. Under parsimony only four independent origins of dimorphism are required: in the theridiid genus Tidarren, in the distal nephilines, in the "argiopoid clade," and in the araneid genus Kaira. Dimorphism has reversed to monomorphism at least seven times, all of them within the large "argiopoid clade." The four independent origins of dimorphism represent two separate instances of an increase in female size coupled with a decrease of male size (involving only two genera), and two separate instances of an increase in female size with male size either remaining the same or increasing, but not as much as females (involving 30 genera). In orb-weaving spiders, far more taxa are sexually dimorphic as a result of female size increase (22 genera) than as a result of male size decrease (two genera). SSD in orb-weaving spiders encompasses several independent evolutionary histories that together suggest a variety of evolutionary pathways. This multiplicity strongly refutes all efforts thus far to find a general explanation for either the origin or maintenance (or both) of SSD, because the different pathways very likely will require distinctly different, possibly unique, explanations. Each pattern must be understood historically before its origin and maintenance can be explained in ecological and evolutionary terms. The most frequently cited example of male dwarfism in spiders, the golden orb-weaving spider genus Nephila (Tetragnathidae), is in fact a case of female giantism, not male dwarfism.     

姜科植物地理

本文讨论了姜科的分类系统、起源、进化和地理分布．姜科为一还热带分布科,按Ｂｕｒｔｔ［８］的系统分２亚科４族．全世界有５２属,约１３７７种,其中姜亚科含４８属,１２６８种．主要分布于热带亚洲．其现代分布中心在印度－马来西亚。闭鞘姜亚科含４属,１０９种,主要分布于热带美洲及非洲。本文在化石资料及现代分布资料的基础上,讨论了姜科的早期分化时间、地点及现代分布格局形成。化石记录表明．欧洲、北美及印度的白垩纪、早第三纪均发现过姜科的化石,据此姜科植物的起源时间应不晚于早白垩纪。姜亚科的早期分化中心推论在劳亚古陆的南部．欧洲和北美没有现代姜科的分布是因为第三纪冰期的影响．而亚洲热带地区现代姜科植物繁盛是因为气候适宜．且相对稳定所致．南美的姜亚科种类应是由非洲传人．而大洋洲的姜亚科种类则是由马来西亚传入．闭鞘姜亚科的早期分化中心推论在西冈瓦纳古陆．亚洲及大洋洲的闭鞘姜亚科的种类应是随印度板块飘向亚洲时传入。中国姜科植物有２２属．２０９种（占全世界属的４２％．种的１５％）．主要分布于马来西亚亚区（占全国属的９０％）．其次为中国喜马拉雅亚区（占全国属的６８％）。最少为中国－日本亚区（占全国属的４５％）。统计数字表明．马来西亚     

Multiple origin of the tropical forest tree family Icacinaceae

Analyses of DNA sequences from four genes (ndhF, rbcL, atpB, and 18S rDNA) and morphological data show that the members of the tropical forest tree family Icacinaceae do not have a common origin. All of the genera earlier placed in Icacinaceae are euasterids but placed in the following three different orders: Garryales, Aquifoliales, and Apiales. Icacina and related genera are members of Garryales and, pending more data, are still best treated as Icacinaceae (sensu stricto). The genera related to Aquifoliales are placed in Cardiopteridaceae and a new family, Stemonuraceae. The genus Pennantia is a member of Apiales and the family Pennantiaceae is recognized. Morphological characters delimiting these groups are discussed. Twenty-six new ndhF sequences were obtained for the study (25 from former Icacinaceae and 1 from Cardiopteris).     

楝科（Meliaceae）的地理分布

楝科为泛热带分布科,全世界有５１属,约５５０—６００种,分布于旧世界热带地区有４６属,热带美洲有８属．热带亚洲和热带非洲为楝科两大现代分布中心．中国楝科共１５属,６１种,占世界属总数的２９％,种总数的１０％。中国楝科的分布是在全球楝科分布区的边缘,主要分布于中国西南部及南部诸省,种类由西南向东南递减。中国楝科属的分布区类型可归为５类：１．热带亚洲、非洲和中南美洲间断分布（１属）;２．旧世界热带分布（３属）;３,热带亚洲至热带大洋洲分布（２属）;４．热带亚洲至热带非洲分布（１属）;５．热带亚洲分布（８属）。中国楝科种的分布区类型仅有２类：１．热带亚洲分布（３１种）;２．中国特有分布（３０种）。楝科植物的起源推断在早白垩纪。中国楝科植物由印度—马来西亚成分及特有成分组成。热带亚洲的楝科植物主要是通过中南半岛和中国云南。广西和海南等地发生联系,而菲律宾和台湾之间可有直接的联系。     

TRADITIONAL GENERIC CONCEPTS VERSUS 18S rRNA GENE PHYLOGENY IN THE GREEN ALGAL FAMILY SELENASTRACEAE (CHLOROPHYCEAE, CHLOROPHYTA)

Coccoid green algae of the Selenastraceae were investigated by means of light microscopy, TEM, and 18S rRNA analyses to evaluate the generic concept in this family. Phylogenetic trees inferred from the 18S rRNA gene sequences showed that the studied species of autosporic Selenastraceae formed a well-resolved monophyletic clade within the DO group of Chlorophyceae. Several morphological characteristics that are traditionally used as generic features were investigated, especially the arrangement of autospores in the mother cells, colony formation, and pyrenoid structure. The parallel arrangement of autospores was confirmed for the genera Ankistrodesmus , Podohedriella , and Quadrigula. In mother cells of Monoraphidium and Kirchneriella , the autospores were arranged serially. Colony formation was either stable ( Quadrigula ) or variable ( Ankistrodesmus , Podohedriella ) within genera. All strains studied possessed naked or starch-covered pyrenoids within the chloroplast. The pyrenoid matrix was homogenous or penetrated by thylakoids. In contrast to considerations of traditional systematics, the present study showed that the presence and structure of pyrenoids are unsuitable for differentiation of genera in Selenastraceae. Furthermore, the molecular analyses showed that any morphological criterion considered so far is not significant for the systematics of the Selenastraceae on the generic level. Species assigned to different genera such as Ankistrodesmus and Monoraphidium were not monophyletic and therefore not distinguishable as separate genera. Species of Monoraphidium appeared in four different lineages of the Selenastraceae. Our phylogenetic analyses support earlier discussions to abandon the common practice of conceiving "small" genera (i.e. genera that are differentiated from other genera by only a few diacritic characteristics and that contain only a small number of species) and to reestablish "large" genera of Selenastraceae such as Ankistrodesmus.     

Evidence for a common evolutionary origin of coronavirus spike protein receptor-binding subunits

Among different coronavirus genera, the receptor-binding S1 subunits of their spike proteins differ in primary, secondary, and tertiary structures. This study identified shared structural topologies (connectivity of secondary structural elements) in S1 domains of different coronavirus genera. The results suggest that coronavirus S1 subunits share a common evolutionary origin but have attained diverse sequences and structures following extensive divergent evolution. The results also increase understanding of the structures and functions of coronavirus S1 domains whose tertiary structures are currently unknown.     

Comparative studies of vegetative anatomy in the Theaeeae of Sri Lanka

Anatomical characters of the mature leaf blade, petiole, young stem and wood were examined in order to substantiate taxonomic boundaries between genera and species of Theaeeae in Sri Lanka. Two species of Temstroemia , one of Adinandra , four of Eurya and two of Gordonia were studied.
The presence of sclereids in most organs of the plant was found to be a common character within the family. However, three distinct types of sclereid are reported. Two basic types of stomata, anomocytic and gordoniaceous, are recognized. The origin of phellogen in the young stems is pericyclic in Gordonia and subepidermal in the other three genera. The species within these genera also appear to have several distinctive characters of their own. Important wood characters have been tabulated in order to illustrate the primitive and advanced characters. Gordonia , especially, exhibits anatomical features which are quite different from those of Temstroemia, Adinandra send Eurya.     

Molecular Evolution of Mollusc Shell Proteins: Insights from Proteomic Analysis of the Edible Mussel Mytilus

Shell matrix proteins (SMPs) that are embedded within calcified layers of mollusc shells are believed to play an essential role in controlling the biomineral synthesis and in increasing its mechanical properties. Among the wide diversity of mollusc shell textures, nacro-prismatic shells represent a tremendous opportunity for the investigation of the SMP evolution. Indeed, nacro-prismatic texture appears early in Cambrian molluscs and is still present in the shell of some bivalves, gastropods, cephalopods and very likely also, of some monoplacophorans. One key question is to know whether these shells are constructed from similar matrix protein assemblages, i.e. whether they share a common origin. Most of the molecular data published so far are restricted to two genera, the bivalve Pinctada and the gastropod Haliotis. The shell protein content of these two genera are clearly different, suggesting independent origins or considerable genetic drift from a common ancestor. In order to describe putatively conserved mollusc shell proteins, here we have investigated the SMP set of a new bivalve model belonging to another genera, the edible mussel Mytilus, using an up-to-date proteomic approach based on the interrogation of more than 70,000 EST sequences, recently available from NCBI public databases. We describe nine novel SMPs, among which three are completely novel, four are homologues of Pinctada SMPs and two are very likely homologues of Haliotis SMPs. This latter result constitutes the first report of conserved SMPs between bivalves and gastropods. More generally, our data suggest that mollusc SMP set may follow a mosaic pattern within the different mollusc models (Mytilus, Pinctada, Haliotis). We discuss the function of such proteins in calcifying matrices, the molecular evolution of SMP genes and the origin of mollusc nacro-prismatic SMPs.     

杜鹃花科白珠树属分子系统学和生物地理学研究进展

白珠树属(Gaultheria) 在杜鹃花科(Ericaceae)系统演化中占有十分重要的地位, 其系统位置和演化关系一直备受争议。最近的分子系统学研究认为, 白珠树属已经不再属于传统上的越橘亚科(Vaccinioideae)的綟木族(Andromedeae), 而是与一些相关属组成了白珠树族(Gaultherieae)。对白珠树属产于美洲的类群和相关类群的分子系统学的初步研究则表明, 该属与Diplycosia、Tepuia和Pernettya等属(均为“常绿类群”)关系密切, 可能应将这几个属并入到白珠树属中, 但其属下分类系统关系还需要对产于亚洲的类群进行深入的研究后才能确定。白珠树属与其近缘属的进化历史和生物地理学关系较为复杂, 与杜鹃花科其他大多数属不同, 白珠树属为典型的环太平洋分布。关于白珠树属的起源问题存在两种不同的推测: 一种观点认为该属起源于南半球的冈瓦纳古陆; 另一种观点则认为其起源于北半球的劳亚古大陆。本文概述了近年来白珠树属的分子系统学和生物地理学研究进展, 并对该属尚存的一些问题进行了讨论。     

Origin and differentiation of endemism in the flora of China

The present paper analyzed 239 endemic genera in 67 families in the flora of seed plants in China.The results showed that there are five families containing more than ten endemic genera,namely,Gesneriaceae (27),which hereafter refers to the number of endemic genera in China,Composite (20),Labiatae (12),Cruciferae (11),and Umbelliferae (10),15 families with two endemic genera,and another 30 families with only one endemic genus.Four monotypic families (Ginkgoaceae,Davidiaceae,Eucommiaceae and Acanthochlamydaceae)are the most ancient,relict and characteristic in the flora of seed plants in China.Based on integrative data of systematics,fossil history,and morphological and molecular evidence of these genera,their origin,evolution and relationships were discussed.In gymnosperms,all endemic genera are relicts of the Arctic-Tertiary flora,having earlier evolutionary history,and can be traced back to the Cretaceous or to the Jurassic and even earlier.In angiosperms,the endemic genera are mostly relicts,and are represented in all lineages in the"Eight-Class System ofClassification of Angiosperms",and endemism can be found in almost every evolutionary stage of extant angiosperms.The relict genera once occupied huge areas in the northern hemisphere in the Tertiary or the late Cretaceous,while neo-endemism mostly originated in the late Tertiary.They came from Arctic-Tertiary,Paleo-tropical-Tertiary and Tethys-Tertiary florisitic elements,and the blend of the three elements with many genera of autochthonous origin.The endemism was formed when some dispersal routes such as the North Atlantic Land Bridge,and the Bering Bridge became discontinuous during the Tertiary,as well as the climate change and glaciations in the late Tertiary and the Quaternary.Therefore,the late Tertiary is the starting point of extant endemism of the flora in China.     

Molecular phylogeny of the volvocine flagellates.

Phylogenetic studies of approximately 2,000 bases of sequence from the large and small nuclear-encoded ribosomal RNAs are used to investigate the origins of the genus Volvox. The colonial and multicellular genera currently placed in the family Volvocaceae form a monophyletic group that is significantly closer phylogenetically to Chlamydomonas reinhardtii than it is to the other unicellular green flagellates that were tested, including Chlamydomonas eugametos, Chlorella pyrenoidosa, and Haematococcus lacustris. Statistical analysis of 251 phylogenetically informative nucleotide positions rejects the "volvocine lineage" hypothesis, which postulates a monophyletic evolutionary progression from unicellular organisms (such as Chlamydomonas), through colonial organisms (e.g., Gonium, Pandorina, Eudorina, and Pleodorina) demonstrating increasing size, cell number, and tendency toward cellular differentiation, to multicellular organisms having fully differentiated somatic and reproductive cells (in the genus Volvox). The genus Volvox appears not to be monophyletic. Volvox capensis falls outside a lineage containing other representatives of Volvox (V. aureus, V. carteri, and V. obversus), and both of these Volvox lineages are more closely related to certain colonial genera than they are to each other. This implies either a diphyletic origin of Volvox from different colonial volvocacean ancestors, a phylogenetic derivation of some of the colonial genera from a multicellular (i.e., Volvox) ancestor, or both. Considered together with previously published observations, these results suggest that the different levels of organizational and developmental complexity found in the Volvocaceae represent alternative stable states, among which evolutionary transitions have occurred several times during the phylogenetic history of this group.     

Chromosome patterns and nuclear phenomena in the cycad families Stangeriaceae and Zamiaceae

The chromosome karyotypes are described and scale diagrams prepared of 35 species representing 8 genera of the cycad families Stangeriaceae and Zamiaceae. The karyotype patterns, chromosome types and characteristics of the nucleolar organisers and heterochromatin are discussed in relation to their evolution in this ancient group of plants. A possible path of chromosome evolution is suggested by the concurrence in some genera of terminal heterochromatic knobs on chromosome arms and a single pair of telocentric homologues which in the remaining genera are replaced by complex partially heterochromatic nucleolar-organising chromosomes. — It is suggested that telocentric chromosomes may be of ancient origin in the cycads and that there may have been a progression from telocentric to mesocentric karyotypes. The paucity of genera today and the taxonomic isolation of the cycads from the remainder of the present-day flora renders impossible the corroboration of this theory by direct cytological comparisons with other groups.     

Floristic relationships among vegetation types of new zealand and the southern andes: similarities and biogeographic implications

Background and Aims

Similarities between the floras of geographically comparable regions of New Zealand (NZ) and the southern Andes (SA) have interested biologists for over 150 years. The present work selects vegetation types that are physiognomically similar between the two regions, compares their floristic composition, assesses the environmental factors that characterize these matching vegetation types, and determines whether phylogenetic groups of ancestral versus modern origin are represented in different proportions in their floras, in the context of their biogeographic history.

Methods

Floristic relationships based on 369 genera of ten vegetation types present in both regions were investigated with correspondence analysis (CA) and ascending hierarchical clustering (AHC). The resulting ordination and classification were related to the environmental characteristics of the different vegetation types. The proportions of different phylogenetic groups between the regions (NZ, SA) were also compared, and between forest and non-forest communities.

Key Results

Floristic similarities between NZ and SA tend to increase from forest to non-forest vegetation, and are highest in coastal vegetation and bog. The floras of NZ and SA also differ in their phylogenetic origin, NZ being characterized by an ‘excess’ of genera of basal origin, especially in forests.

Conclusions

The relatively low similarities between forests of SA and NZ are related to the former being largely of in situ South American and Gondwanan origin, whereas the latter have been mostly reconstituted though transoceanic dispersal of propagules since the Oligocene. The greater similarities among non-forest plant communities of the two regions result from varied dispersal routes, including relatively recent transoceanic dispersal for coastal vegetation, possible dispersal via a still-vegetated Antarctica especially for bog plants, and independent immigration from Northern Hemisphere sources for many genera of alpine vegetation and grassland.Key words: Biogeographic history, floristic similarities, generic composition, local floras, New Zealand, phylogenetic origin, southern Andes, vegetation types     

Generic names in the Orbiliaceae (Orbiliomycetes) and recommendations on which names should be protected or suppressed

A list of all generic names that have been connected with the Orbiliomycetes is provided. Recommendations are made as to which names should be used in accordance with the rules and the different generic concepts. There is a mismatch in the current generic concepts within Orbiliomycetes regarding the two morphs: a narrow concept is used for the asexual morphs, but a broad concept relies on the sexual morphs. As a consequence, many more generic names have been established for the asexual morphs. A number of previous generic concepts are artificial, since they were based on single characters without molecular support. In order to provide solutions for this mismatch, we present three different generic concepts within the Orbiliomycetes. A broad concept recognizes a large genus Orbilia, with which most of the listed names fall into synonymy, but could be maintained as infrageneric names. Due to the lack of data proving phylogenetic relationships, this broad concept, at present, is the most practicable and recommended one. A moderate concept subdivides Orbilia into several genera, with all nematode-trapping fungi merged in Arthrobotrys. A narrow generic concept accepts genera based on differences in trapping organs, but also subdivides the remaining groups of Orbilia into additional genera. Trapping of invertebrates (zoophagy) is not restricted to Arthrobotrys in a broad sense, but occurs also in the more distant basal genera Hyalorbilia and Lecophagus, which mainly prey on rhizopods and rotifers. Whether these predatory capabilities trace back to a common ancestor is not clear. The following new combinations are proposed: Hyalorbilia oviparasitica, Hyalorbilia quadridens, Hyalorbilia tenuifusaria, and Orbilia fissilis.     

Discontinuous DNA variation in the evolution of plant species

The four genera,Clarkia, Nicotiana, Lathyrus andAllium have widely different geographical distributions and evolutionary patterns, but the nucleardna amounts of species within each genus occur in groups at intervals of approximately 2 or 4 picograms. Diploids with different basic chromosome numbers, and polyploids, can occur in the same group. When anydna group of one genus is paired with the nearestdna group of another genus all other groups of both genera pair exactly in so far as they have a common range of DNA amounts. Hence thedna groups of the four genera are associated in larger groups, or nodes, separated by 2 or 4 picograms forming a progression over all four genera, implying there are favoured, discontinuous,dna amounts, either arisingde novo or subsequently within a continuously varying distribution.     

Correlations between flavonoid chemistry,anatomy and geography in the restionaceae

Comparisons of the flavonoid patterns in stems and inflorescences between Australasian and South African members of the Restionaceae indicate significant differences with geography. Nine of 14 Australasian species contain gossypetin or a related 8-hydroxyflavonoid and proanthocyanidins are uncommon. By contrast, the 33 South African taxa studied contain common flavonols, flavones and glycoflavones, while proanthocyanidins are present in 29. Two anatomically related South African genera, Chondropetalum and Elegia, contain, in addition, myricetin 3-galactoside, together with the 3-galactosides of the myricetin methyl ethers, larycitrin and syringetin. These results confirm the conclusions derived from anatomy that members of Hypolaena, Leptocarpus and Restio, genera represented in both Australia and South Africa, have the distinctive flavonoids characteristic of their geographic origin rather than of their systematic position. The family as a whole is different in flavonoid pattern from other monocotyledonous families with which it is sometimes associated.     

亚洲中部荒漠区的植物特有属

系统地列出了亚洲中部荒漠区的13个植物特有属或近特有属——绵刺属、沙冬青属、四合木属、百花篙属、革苞菊属、河西菊属、喀什菊属、紊篙属、连蕊芥属、钝基草属、合头黎属、戈壁黎属、霸王属。这些属都是单种属或双种属,分类上属于孤立的类群;演化系统上是既起源古老而又进化的类群;生态上都是早生植物,绝大多数是荒漠种,其生活型灌木和半灌木占优势;地理分布上阿拉善荒漠区是其特有属的分布中心;起源上是多元的。     

Macromutation and evolution: the fixation of Goldschmidt's macromutations as species and genus traits. Hairlessness mutations in mammals

A brief survey of the development of concepts on the role of macromutations in evolution is given. Contrary to Iu. A. Filipchenko (1926, 1927), who introduced the "micro- and macromutation" terms and believed that regularities of macroevolution could not be reduced to microevolutionary processes, the majority of "synthetists" explained any form of evolution by changes in allele frequencies. From the studies of Drosophila homoeotic mutants R. Goldschmidt (1940) developed the concept of "hopeful monsters" and their role in macroevolution. However, the homoeotic mutants are of drastically reduced viability, which allows the gradualists to reject Goldschmidt's ideas. The distribution of hairlessness mutations (hairless, nude etc.) with the monogenic pattern of inheritance in mammals was studied. Hairless mutants are known in Peromyscus, Mus musculus, Rattus rattus, R. norvegicus, Canis familiaris, Ovis aries. Hairlessness as norm is found in 53 among contemporaneous 1037 mammalian genera. Part of these cases (hairlessness in all Cetacea and Sirenia) may be explained in terms of both macromutations and obligatory gradualism. There is no doubt as to the macromutational origin of hairlessness in the bat Cheiromeles and the rodent Heterocephalus (Bathyergidae); the genera systematically and ecologically close to these have normal pelage. It is quite possible that hairlessness of walrus (Odobenus) has the same origin. The appearance and fixation of single Goldschmidt's macromutation cannot yet be considered as a macroevolutionary process, though the possibility of fixation of a macromutation in nature as a species and genus character contradicts strongly the concept of obligatory gradualism of evolution.     

Molecular identification of sequestered diatom chloroplasts and kleptoplastidy in foraminifera

Kleptoplastidy is the ability of heterotrophic organisms to preserve chloroplasts of algal preys they eat and partially digest. As the sequestered chloroplasts stay functional for months, the "host" becomes photosynthetically active. Although remaining a marginal process, kleptoplastidy was observed in different protist lineages, including foraminifera. Previous studies showed at least eight species of the foraminiferal genera Haynesina and Elphidium grazing on diatoms and husbanding their chloroplasts. In order to characterize more precisely the origin of kleptochloroplasts in these genera, we obtained 1027 chloroplastic 16S rDNA sequences from 13 specimens of two Haynesina and five Elphidium species. We identified the foraminiferal kleptochloroplasts using a reference phylogeny made of 87 chloroplastic sequences of known species of diatoms and brown algae. All the analyzed specimens were performing kleptoplastidy and according to our phylogenetic analyses they seem to retain exclusively chloroplasts of diatom origin. There is no apparent specificity for the type of diatom from which chloroplasts originated, however some foraminiferal species seem to accept a wider range of diatoms than others. Possibly the diversity of kleptochloroplasts depends on the type of diatoms the foraminiferans feed on.     

Paleoclimatic history and vicariant speciation in the "sand goby" group (Gobiidae, Teleostei)

Vicariant and climatic cycling speciation hypotheses of the 'sand gobies' belonging to the genera Pomatoschistus, Gobiusculus, Knipowitschia, and Economidichthys are tested using molecular phylogenies constructed of nuclear DNA (ITS1 locus) and mitochondrial DNA (12S and 16S fragments). These gobies are among the most abundant in the Eastern Atlantic-Mediterranean region, and play an important role in the ecosystem. Considerable ITS1 length differences, primarily due to the presence of several tandem repeats, were found between species and even within individuals. Therefore, phylogenetic analyses focused on fragments of the 12S and 16S mtDNA region that have been sequenced for 16 goby taxa. The 'sand gobies' clustered as a monophyletic group as proposed on morphological grounds. However, G. flavescens, E. pygmaeus, and K. punctatissima clustered within the Pomatoschistus species, pointing to a paraphyletic origin of these genera. Furthermore, the genetic divergence between P. minutus from the Adriatic Sea versus the Atlantic-Mediterranean region was as high as the divergence within the P. minutus complex, suggesting that P. minutus from the Adriatic Sea should be considered as a distinct species. The "star" phylogeny might suggest that these gobies evolved in a very short time period, possibly linked to the drastic alterations in the Mediterranean Sea during and immediately after the Messinian salinity crisis at the end of the Miocene. The freshwater life-style appeared monophyletic; equating its origin with the salinity crisis resulted in a molecular clock estimate of 1.4% divergence per million years. The last common ancestor probably occupied sandy bottoms and a coastal niche while several species subsequently adapted to new habitats (pelagic, freshwater or stenohaline). The origin of the shallowest clades dated back to the glacial cycling during the Pleistocene epoch.