Ontogenetic variability in the external morphology of monkey goby, Neogobius fluviatilis (Pallas, 1814) and its relevance to invasion potential

In the previous decade, four species of non-native gobies have invaded the middle section of the river Danube and its tributaries. An effective tool for understanding biological invasions is the evaluation of various biological traits (morphological, life history, ontogenetic) within an epigenetic context. The present study examines the external morphology of monkey goby Neogobius fluviatilis (Pallas, 1814) from the mouth of the River Hron, the morphological differences among three goby species (monkey, bighead and round) and the relevance of these differences for invasive potential. Monkey goby reach their definite phenotype very early in their ontogeny and thus represent a strongly precocial (specialized) species with direct development. The morphological differences between monkey and two other goby species also reflect its strong specialization for sandy substrata and smaller prey types. Thus, monkey goby are not expected to spread to new areas as fast as the round and bighead gobies, and their distribution is likely to be limited to sandy and/or sandy-gravel substrata. If this assumption is correct, then the potential adverse impact of monkey goby on native fauna or even ecosystem is likely to be less than that of the bighead and round gobies. Handling editor: K. Martens     

Feeding preferences of an invasive Ponto‐Caspian goby for native and non‐native gammarid prey

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Comparative energy allocation in two sympatric, closely related gobies: the black goby Gobius niger and the grass goby Zosterisessor ophiocephalus

Seasonal energy allocation of lipid reserves into different body tissues was analysed comparatively in two sympatric, closely related gobies: the grass goby Zosterisessor ophiocephalus and the black goby Gobius niger . Lipid reserves were measured in liver, muscle and ovary and compared between the two species within a given sex and seasonal period (reproductive v . non-reproductive). Furthermore, temporal patterns of lipid reserves were investigated in the two species in relation to gonado-somatic and liver-somatic indices, as well as the relationship between size and lipid content. Results showed that the grass goby allocated more lipid reserves in reproduction while the black goby accumulated more reserves in liver and muscle, at a given size, although the temporal patterns of lipid accumulation and depletion were basically similar. Results are discussed in the light of life-history theories, taking into account both adaptation and evolutionary constraints.     

Distribution,abundance and condition of invasive Ponto‐Caspian gobies Ponticola kessleri (Günther, 1861), Neogobius fluviatilis (Pallas, 1814), and Neogobius melanostomus (Pallas, 1814) in the Sava River basin,Croatia

Distributions, population densities, invasive potentials and adverse impacts of invasive Ponto‐Caspian gobies on native fauna in the Sava River basin remain unknown, whereby 23 locations in the Sava basin were sampled during a three‐year period (2011–2013). Among the five Ponto‐Caspian goby species previously reported in Croatia, only Neogobius fluviatilis (291), Neogobius melanostomus (177) and Ponticola kessleri (21) specimens were collected. Proterorhinus semilunaris and Babka gymnotrachelus were not found. N. fluviatilis was dominant among the invasive gobies at the majority of locations and almost omnipresent (apart from the upper Sava reaches) in the Sava basin. N. melanostomus was found only in the navigable Sava reaches, but its distribution range still seems to be expanding. The status of P. kessleri invasion remains unclear, as it was only found in 2011 at three locations in the lower Sava reaches. No significant (P > 0.05) increase in average catch‐per‐unit‐effort (CPUE) values was observed during the three‐year period. There is some evidence that a decline in Gobio gobio populations might be expected in response to increasing N. fluviatilis population densities. The findings suggest that further range expansions by the Ponto‐Caspian gobies can be expected in Croatia and the implementation of measures to limit their spreading should be considered.     

The Primary Lower Gypsum in the Mediterranean: A new facies interpretation for the first stage of the Messinian salinity crisis

The detailed facies and physical stratigraphic analysis of the Primary Lower Gypsum in the Mediterranean indicates a surprising bed-by-bed correlation at basin-scale (Spain, Italy, Hellenic arc and Cyprus arc), that is tuned to the orbital calibration for the first stage of the Messinian salinity crisis from 5.96 to 5.61 Ma ago. A total of 16, precessionally-controlled, gypsum cycles were deposited rapidly in less than 350 ka, forming sequences up to 300 m thick. The lack of subaerial exposure features and the common facies associations and stacking pattern for sections located thousands of kilometers apart in different geological settings indicates a modest depositional depth, not extremely shallow. Selenite deposition occurred only at the bottom of restricted marginal basins less than 200 m deep, while no gypsum could precipitate in the deeper euxinic Mediterranean portions where only thin and barren shale/dolostone couplets formed. The lowermost selenite beds pass laterally to dolomite-rich limestones interbedded with barren euxinic shales in poorly oxygenated settings, indicating that the gypsum sedimentation was diachronous and did not necessarily mark the onset on the Messinian salinity crisis.Evaporite facies sequences (EF1 to 8) within individual gypsum beds show small-scale, subaqueous sedimentary cycles that mimic regressive-transgressive cycles: a) initial evaporite precipitation at relatively low supersaturation produced the massive selenite (facies EF3) in a relatively deep setting; b) continuous evaporation and drawdown by oscillating brine level formed the banded selenite (EF4) at the aridity acme of the precessionally-controlled cycle; c) general progressive brine level rise with strong brine flow led to the formation of large selenite supercones branching laterally (supercones in Spain and branching selenite, EF5, in the rest of the Mediterranean); and d) flooding by undersaturated continental water terminated gypsum precipitation with the deposition of argillaceous sediments (EF1, Northern Apennines), and/or limestone (EF2, Sicily and Spain) during the humid phase in the precession climate cycle.The stacking pattern and selenite facies associations suggest an overall shallowing-upward trend with a basin-wide hydrologic change starting from the 6th cycle (5.84 Ma), which is marked by the appearance of the branching selenite facies (supercones) in Spain and indicates that the brines became current-dominated. The Sr-isotope stratigraphy suggests that in the Northern Apennines the brines were strongly modified by continental waters (⁸⁷Sr/⁸⁶Sr = 0.708893 to 0.708998), and received direct pulses of Atlantic seawater (⁸⁷Sr/⁸⁶Sr = 0.70900 to 0.709024) only in the upper part of the section. In areas away from the mainland, such as Sicily, the continental input was by far less important.     

Migration and isolation during the turbulent Ponto‐Caspian Pleistocene create high diversity in the crustacean Paramysis lacustris

The Ponto‐Caspian brackish‐water fauna inhabits estuaries and rivers of the Black, Azov and Caspian seas and is fragmented by higher salinity waters and a major interbasin watershed. The fauna is known for the high levels of endemism, complex zoogeographic histories, and as a recent source of successful invasive species. It remains debated whether the Black and Azov Sea brackish‐water populations survived unfavourable Pleistocene conditions in multiple separate refugia or whether the two seas were (repeatedly) recolonized from the Caspian. Using microsatellite and mtDNA markers, we demonstrate deep among‐ and within‐basin subdivisions in a widespread Ponto‐Caspian mysid crustacean Paramysis lacustris. Five genetic clusters were identified, but their relationships did not reflect the geography of the region. The Azov cluster was the most distinct (4–5% COI divergence), despite its geographic position in the corridor between Black and Caspian seas, and may represent a new species. In the northern Black Sea area, the Dnieper cluster was closer to the Caspian cluster than to the neighbouring Danube–Dniester–Bug populations, suggesting separate colonizations of the Black Sea. Overall, the data implied a predominant gene flow from the east to the Black Sea and highlight the importance of Caspian Sea transgressions in facilitating dispersal. Yet, the presence of distinct lineages in the Black Sea points to the persistence of isolated refugial populations that have gained diagnostic differences under presumably high mutation rates and large population sizes. The unfavourable Pleistocene periods in the Black Sea therefore appear to have promoted diversification of the brackish‐water lineages, rather than extirpated them.     

Ten polymorphic microsatellite markers in the invasive round goby (Neogobius melanostomus) and cross‐species amplification

Ten tetranucleotide, dinucleotide and compound microsatellite loci were isolated and characterized for the round goby, Neogobius melanostomus. Analysis of 64 gobies from one nonindigenous population in Lake Erie, Ontario, Canada, indicated that allele number varied from three to 12 per locus, while observed heterozygosity ranged between 0.33 and 0.86. Eight of these primers showed some amplification in other species in four genera. These newly developed microsatellite markers are a powerful tool that will provide insights into population structure and dispersal of the round goby in their novel environment.     

Sound production by the Shi drum Umbrina cirrosa and comparison with the brown meagre Sciaena umbra: a passive acoustic monitoring perspective

Sounds produced by the Shi drum Umbrina cirrosa were short trains of pulses with an average pulse period of 180 ms, pulse duration of c. 40 ms and an average peak frequency of 400 Hz; average values of acoustical properties differed from those recorded from the brown meagre Sciaena umbra in previous studies. The present study provides a preliminary tool for discriminating between these two species while conducting passive acoustic monitoring. The potential effects of ontogeny on sound production in both species are discussed and recommendations are made for further research.     

Species Composition and Distribution of Invasive Ponto‐Caspian Amphipods in the Off‐Channel Microhabitats of a Temperate,Lowland Dam Reservoir

The Włocławek Dam Reservoir located on the lower Vistula River (central Poland) is part of the central corridor used by Ponto‐Caspian species to migrate in Europe. It provides a number of habitats suitable for sustaining populations of several non‐indigenous taxa. Four Ponto‐Caspian amphipod species were recorded in the reservoir: Chaetogammarus ischnus, Chelicorophium curvispinum, Dikerogammarus haemobaphes and Pontogammarus robustoides. We found significant differences in amphipod densities and species composition among various microhabitats in off‐channel areas. P. robustoides was the only amphipod species that occurred on very shallow (<1 m) sandy bottom near the shore. It inhabited also other sites but its share in the total number of amphipods and abundance decreased with the distance from the shore. Furthermore, at sites more distant from shore its affinity for plant substratum was higher. D. haemobaphes, C. ischnus and C. curvispinum clearly preferred sites distant from shore, overgrown by macrophytes or covered by mussel shells. Furthermore, the abundance of C. ischnus was negatively correlated with the presence of P. robustoides. (© 2008 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Molecular markers for systematic identification and population genetics of the invasive Ponto‐Caspian freshwater gammarid Dikerogammarus villosus (Crustacea,Amphipoda)

The Ponto‐Caspian amphipod, Dikerogammarus villosus, is an invasive species of many European rivers. First, we show that size difference of nrDNA ITS1 allows discriminating D. villosus from Dikerogammarus bispinosus, a closely related but morphologically hardly distinguishable species. Second, we present two types of polymorphic markers for D. villosus, three microsatellites and two single nucleotide polymorphisms (SNPs) of mtDNA COI gene, which were scored by polymerase chain reaction‐single strand conformational polymorphism (PCR‐SSCP). These markers will be very useful in studying population genetics of D. villosus.     

Isolation of novel microsatellite loci in the black goby Gobius niger and cross‐amplification in other gobiid species (Perciformes,Gobiidae)

Twelve microsatellite loci were isolated from and characterized for the black goby Gobius niger. These loci were tested on a total of 48 individuals from two geographically distant locations (Orbetello and Cattolica) and the number of alleles ranged from two to 18, with expected (H_e) and observed (H_o) heterozygosities ranging from 0·042 to 0·941 and from 0·042 to 0·917, respectively. The loci described were used to cross‐amplify other gobiid species belonging to Gobius, Zosterisessor, Lesueurigobius and Aphia.     

The morphology of the eutherian ethmoidal region and its implications for higher‐order phylogeny

The ethmoidal region is the most rostral part of the mammalian cranium and provides the surface for the olfactory epithelium. It is part of the respiratory tract and comprises certain mechanic and mechanosensoric functions depending on the species’ mode of foraging. It is one of the least studied aspects of cranial anatomy. A complete and precise characterization of the previously undescribed fetal nasal regions of the aardvark (Orycteropus afer, Tubulidentata), the African elephant (Loxodonta africana, Proboscidea) and the rock hyrax (Procavia capensis, Hyracoidea) is given. The morphology of the ethmoidal region in O. afer and L. africana, highly macrosmatic mammals, is complicated and in many aspects derived, whereas the nasal structures in P. capensis are fairly simple. In order to contribute to current debates on higher‐order eutherian systematics 24 skeletal and soft‐tissue characters in 26 mammalian taxa (25 eutherian taxa and 1 marsupial as outgroup) were evaluated and optimized onto recent eutherian phylogenies (one morphology and one molecular based phylogeny, respectively). It is shown that the ethmoidal region is highly variable within eutherians and its structural elements display a strong tendency towards homoplasy. Therefore, this cranial region is not suitable for use in mammalian superordinal systematics except as support for monophyletic Tethytheria.     

Complex controls on ostracod palaeoecology in a shallow coastal brackish‐water lake: implications for palaeosalinity reconstruction

1. The low‐Mg calcite shells of Ostracoda (Crustacea) are often well preserved in the sediments of alkaline lakes. In coastal waters that have undergone large temporal changes in salinity, ostracod assemblages preserved in the sediment record have been used to reconstruct palaeosalinity, often assuming that salinity is the only significant control on the faunas. 2. We evaluate the performance of ostracods as palaeosalinity indicators in Hickling Broad, a shallow brackish coastal lake in Norfolk, U.K., by comparing fossil ostracod assemblages covering two centuries with geochemical inferences and instrumental records of past salinity and water composition along with other palaeolimnological indicators. 3. Despite large changes in the salinity of the lake and the supposed salinity sensitivity of ostracods, the fossil ostracod assemblages do not clearly reflect the salinity trends inferred from the other independent data. Rather, a complex series of changes has occurred in the lake over the past 200 years and factors other than salinity, including eutrophication, toxicity and associated complex alterations in habitat availability have probably influenced ostracod assemblages. In contrast, there is a good broad agreement between inferred or measured salinity and the trace‐element chemistry of ostracod shells. 4. We conclude that ostracod faunas may not always provide unambiguous palaeosalinity records and should therefore not be used to reconstruct salinity changes except as part of a multi‐proxy investigation that includes other palaeoecological and/or geochemical indicators.     

Sex differences in the vocal repertoire of adult red‐capped mangabeys (Cercocebus torquatus): a multi‐level acoustic analysis

Sex differences in the vocal behavior of nonhuman primates can take various forms: sex‐specific call types, differential production of shared call types, or sex discrepancy in phonation. Also, a growing literature is evidencing that systematically analyzing the vocal repertoires of primates at the call level might lead to underestimating their communicative abilities. Here, we present an extensive multi‐level analysis of the still unknown vocal repertoire of adult red‐capped mangabeys (Cercocebus torquatus), with a special emphasis on sex differences. We collected recordings from seven adult males and seven adult females housed in captivity. We present a structurally‐based classification of mangabey calls that we cross‐validated by an analysis of the associated contexts of emission. We found 12 sound units (including six sex‐specific) that were concatenated to form eight call types (including four sex‐specific), which were produced either singularly or in sequences composed of one (“repetition”) or several (“combination”) call types. We extracted organizational principles that ruled call composition and calling patterns. This revealed a high degree of potentially meaningful variability in terms of semantics and syntax. Male–female discrepancy in terms of phonation could be related to morphological dimorphism and would enable listeners to behave appropriately according to the sex of the caller. Sex differences in repertoire size, structural gradation, and call usage could reflect specificities of male–female social roles. We discuss the pertinence of these sex differences according to social system and habitat quality. Am. J. Primatol. 72:360–375, 2010. © 2010 Wiley‐Liss, Inc.     

A molecular phylogeny for the pyraloid moths (Lepidoptera: Pyraloidea) and its implications for higher‐level classification

Pyraloidea, one of the largest superfamilies of Lepidoptera, comprise more than 15 684 described species worldwide, including important pests, biological control agents and experimental models. Understanding of pyraloid phylogeny, the basis for a predictive classification, is currently provisional. We present the most detailed molecular estimate of relationships to date across the subfamilies of Pyraloidea, and assess its concordance with previous morphology‐based hypotheses. We sequenced up to five nuclear genes, totalling 6633 bp, in each of 42 pyraloids spanning both families and 18 of the 21 subfamilies, plus up to 14 additional genes, for a total of 14 826 bp, in 21 of those pyraloids plus all 24 outgroups. Maximum likelihood analyses yield trees that, within Pyraloidea, differ little among datasets and character treatments and are strongly supported at all levels of divergence (83% of nodes with bootstrap ≥80%). Subfamily relationships within Pyralidae, all very strongly supported (>90% bootstrap), differ only slightly from a previous morphological analysis, and can be summarized as Galleriinae + Chrysauginae (Phycitinae (Pyralinae + Epipaschiinae)). The main remaining uncertainty involves Chrysauginae, of which the poorly studied Australian genera may constitute the basal elements of Galleriinae + Chrysauginae or even of Pyralidae. In Crambidae the molecular phylogeny is also strongly supported, but conflicts with most previous hypotheses. Among the newly proposed groupings are a ‘wet‐habitat clade’ comprising Acentropinae + Schoenobiinae + Midilinae, and a provisional ‘mustard oil clade’ containing Glaphyriinae, Evergestinae and Noordinae, in which the majority of described larvae feed on Brassicales. Within this clade a previous synonymy of Dichogaminae with the Glaphyriinae is supported. Evergestinae syn. n. and Noordinae syn. n. are here newly synonymized with Glaphyriinae, which appear to be paraphyletic with respect to both. Pyraustinae and Spilomelinae as sampled here are each monophyletic but form a sister group pair. Wurthiinae n. syn. , comprising the single genus Niphopyralis Hampson, which lives in ant nests, are closely related to, apparently subordinate within, and here newly synonymized with, Spilomelinae syn. n.     

Cyst‐Theca Relationship and Phylogenetic Position of Impagidinium caspienense Incubated from Caspian Sea Surface Sediments: Relation to Gonyaulax baltica and Evidence for Heterospory within Gonyaulacoid Dinoflagellates

We investigate the cyst‐theca relationship of Impagidinium caspienense. Through an incubation experiment, we succeeded in examining the motile stage. Additional molecular analysis of single‐cyst PCR (LSU and SSU rDNA) reveal that the cyst is related to the species Gonyaulax baltica Ellegaard et al. ( 2002 ). The ability of this species to belong to two types of cyst‐based genera (spiniferate and impagidinioid) suggests that environmental (particularly salinity) and not genetic factors explain the formation of both morphotypes by G. baltica, which provides evidence for heterospory in this species. The affiliation to G. baltica demonstrates that I. caspienense is not endemic to the Caspian Sea. The phylogenetic position of several other gonyaulacoid species is also documented: Impagidinium pallidum, Ataxiodinium choane, Pyxidinopsis psilata, Spiniferites belerius, and Spiniferites ramosus. The LSU and SSU rDNA based phylogenies suggest that the genera Impagidinium and Spiniferites are not monophyletic, and that P. psilata and A. choane are close to Gonyaulax verior and Gonyaulax polygramma, respectively. In addition, this study accentuates the importance of cyst morphology in the classification of the Gonyaulacales.     

Isolation and characterization of microsatellites in Neogobius kessleri (Perciformes,Gobiidae) and cross‐species amplification within the family Gobiidae

Fourteen polymorphic microsatellites were isolated from Neogobius kessleri, a benthic fish of Ponto‐Caspian origin which has been recently introduced into the Middle and Upper Danube River. Number of alleles and heterozygosity per locus in a sample of 32 fish individuals ranged from two to four and from 0.13 to 0.75, respectively. These primers will be useful in determining the population structure of N. kessleri. In addition, successful cross‐amplification was obtained for four related species, N. melanostomus, N. fluviatilis, N. gymnotrachelus and Proterorhinus marmoratus. These microsatellite loci may be useful for the evaluation of the origin of non‐native goby populations.     

Cryptic lineage divergence in marine environments: genetic differentiation at multiple spatial and temporal scales in the widespread intertidal goby Gobiosoma bosc

The adaptive radiation of the seven‐spined gobies (Gobiidae: Gobiosomatini) represents a classic example of how ecological specialization and larval retention can drive speciation through local adaptation. However, geographically widespread and phenotypically uniform species also do occur within Gobiosomatini. This lack of phenotypic variation across large geographic areas could be due to recent colonization, widespread gene flow, or stabilizing selection acting across environmental gradients. We use a phylogeographic approach to test these alternative hypotheses in the naked goby Gobiosoma bosc, a widespread and phenotypically invariable intertidal fish found along the Atlantic Coast of North America. Using DNA sequence from 218 individuals sampled at 15 localities, we document marked intraspecific genetic structure in mitochondrial and nuclear genes at three main geographic scales: (i) between Gulf of Mexico and Atlantic Coast, (ii) between the west coast of the Florida peninsula and adjacent Gulf of Mexico across the Apalachicola Bay, and (iii) at local scales of a few hundred kilometers. Clades on either side of Florida diverged about 8 million years ago, whereas some populations along the East Cost show divergent phylogroups that have differentiated within the last 200,000 years. The absence of noticeable phenotypic or ecological differentiation among lineages suggests the role of stabilizing selection on ancestral phenotypes, together with isolation in allopatry due to reduced dispersal and restricted gene flow, as the most likely explanation for their divergence. Haplotype phylogenies and spatial patterns of genetic diversity reveal frequent population bottlenecks followed by rapid population growth, particularly along the Gulf of Mexico. The magnitude of the genetic divergence among intraspecific lineages suggests the existence of cryptic species within Gobiosoma and indicates that modes of speciation can vary among lineages within Gobiidae.     

Diurnal behavioural patterns and spread of the Ponto‐Caspian invader Hemimysis anomala G. O. Sars, 1907 (Crustacea,Mysidacea) in the Elbe River,Czech Republic

The Ponto‐Caspian invader Hemimysis anomala was studied in the river Elbe, Czech Republic. The study confirmed further spread of the species: the presence of juveniles and gravid females indicated that H. anomala is established within the Elbe River. Bottom, mid‐water and surface drift samples were collected during 24‐h cycles from July to September 2007. A tendency to remain hidden was the main behavioural strategy of H. anomala in the riverine environment. The mysid was found mainly in the bottom and mid‐water layers, and observed density and mean length of drifting individuals increased with decreasing light levels, being highest at night. Similarly, density and mean length of drifting individuals increased with increasing turbidity. The effect of turbidity on density was particularly apparent during the night samples, suggesting further advantage to night‐drifting individuals. (© 2012 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Molecular phylogeny of African hinge‐back tortoises (Kinixys): implications for phylogeography and taxonomy (Testudines: Testudinidae)

We examine the phylogeography, phylogeny and taxonomy of hinge‐back tortoises using a comprehensive sampling of all currently recognized Kinixys species and subspecies and sequence data of three mitochondrial DNA fragments (2273 bp: 12S rRNA, ND4 + adjacent DNA coding for tRNAs, cytb) and three nuclear loci (2569 bp: C‐mos, ODC, R35). Combined and individual analyses of the two data sets using Bayesian and Maximum Likelihood methods suggest that the savannah species of Kinixys are paraphyletic with respect to the rainforest species K. homeana and K. erosa, and that the rainforest species may be derived from a savannah‐living ancestor. The previously recognized savannah species K. belliana was a conglomerate of three deeply divergent clades that we treat here as distinct species. We restrict the name K. belliana (Gray, 1830) to hinge‐back tortoises ranging from Angola to Burundi, while five‐clawed hinge‐back tortoises from the northernmost part of the formerly recognized range of K. belliana, together with four‐clawed tortoises from West Africa, are assigned to the species K. nogueyi (Lataste, 1886). These two species are allied to K. spekii, whereas Southeast African and Malagasy hinge‐back tortoises formerly lumped together with K. belliana represent the distinct species K. zombensis Hewitt, 1931, which is sister to K. lobatsiana. The latter two species together constitute the sister group of the rainforest species K. homeana and K. erosa. Mitochondrial data suggest that K. natalensis has a basal phylogenetic position in a clade embracing K. belliana sensu stricto, K. nogueyi and K. spekii, while nuclear data and the two data sets combined favour a sister group relationship of K. natalensis to all other hinge‐back tortoises. Phylogeographic structure is present in all wide‐ranging species and correlates in K. homeana and K. erosa with the Dahomey Gap and former rainforest refugia. The Malagasy population of K. zombensis is weakly differentiated from its South African conspecifics and further sampling is needed to determine whether there is support for the subspecific distinctness of Malagasy tortoises.