Voiceprint identification and its application to sociological studies of wild Japanese monkeys (Macaca fuscata yakui)

Japanese monkeys often exchange the particular vocal sound, “coo,” especially when they feed or move as a group. It was considered that the “coo” sound had no positive social meaning, perhaps because the “coo” sound network and its function were hidden behind other behavioral observations. For identification of the vocalizer only from hearing the “coo” sound, three phonetic values, i.e., the “fundamental,” “duration,” and “formants,” plus other characteristics were used as indices of voiceprints. The results indicated that these were effective for identifying the vocalizer in two-thirds of the adults in the study troop which was composed of 12 adults and 16 immature members. The “coo” sound exchange network among the troop members (adults) was drawn on the basis of the voiceprint identification. The network showed three characteristics as follows: (1) matriarchs of the kin-groups frequently exchanged “coo” sounds with each other; (2) the other females exchanged “coo” sounds mostly within their own kin-groups; and (3) males seldom participated in the “coo” sound exchange. This suggests that “coo” sound exchange plays a central role for the matriarch of kin-groups in binding each kin-group and, ultimately, in binding all members together into an organized troop.     

Forms of output distribution between two individuals motivated by a satisfaction function

Motivations of two individuals governed by a satisfaction function are assumed to determine their respective “efforts”, which result in the production of “output”, i.e., objects of satisfaction. In previous papers the sharing of output was prescribed in advance. In the present article, however, the sharing formula itself is determined to a certain extent by the satisfaction function. The rate of remuneration per unit of output for each individual is taken to be proportional to the derivative of the satisfaction of the other individual with respect to the effort of the first. The formulation of this condition leads to a partial differential equation whose solutions determine the sharing formula. Sharing determined in this way is referred to as sharing according to the Condition of Mutual Need (C.M.N.). Satisfaction resulting from five different situations are the computed and compared: (1) an individual producing and consuming alone; (2) two individuals sharing equally and neither taking the “initiative” to determine the optimum output; (3) sharing determined by C.M.N. with optimum output determined as in (2); (4) equal sharing but with one individual taking “initiative” in determining optimal output; and (5) sharing determined by C.M.N. and optiml output by the “initiative” of one individual. further considerations concern conditions imposed on the arbitrary function occurring in the solution of the above-mentioned partial differential equation.     

On reinforcement and interference between stimuli

It is shown that the current “two-factor” theory of nerve excitation can account for sustained inhibition or enhancement by a sequence of stimulus pulses, and for the decrease in the reinforcement period with each successive pulse of the train.     

Foliage consumption and larval development of parasitized and unparasitized soybean looper,Pseudoplusia includens [Lep.: Noctuidae], reared on a resistant soybean genotype and effects on an associated parasitoid,Compidosoma truncatellum [Hym.: Encyrtidae]

Field grown foliage from the resistant soybean [Glycine max (L.) Merrill] breeding line GAT “81–327” and the susceptible cultivar “Ransom” was used to rear unparasitized larvae of the soybean looper (SBL),Pseudoplusia includens (Walker), and larvae parasitized byCopidosoma truncatellum (Dalman). SBL larvae, whether parasitized or not, consumed more foliage when fed “Ransom”. Unparasitized larvae reared on “81–327” had longer developmental times and suffered greater mortality than unparasitized larvae reared on “Ransom”. Parasitization of SBL larvae byC. truncatellum increased total foliage consumption of both soybean lines. Parasitized larvae reared on the resistant “81–327” weighed less and yielded fewer parasitoid adults.

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Résumé Des larves dePseudoplusia includens (Walker) parasitées ou non parCopidosoma truncatellum (Dalman) ont été nourries des feuilles de deux lignées de soja [Glycine max (L.) Merrill] l'une, GAT “81–327” résistante et l'autre, “Ransom” sensible. Les larves deP. includens qu'elles soient parasitées ou non consommaient plus de feuillage lorsqu'elles étaient nourries de la lignée “Ransom”. Les larves non parasitées élevées sur “81–327” avaient un cycle de développement beaucoup plus long et un taux de mortalité beaucoup plus élevé que les larves non parasitées élevées comparativement sur feuilles de “Ransom”. Par contre, les larves parasitées manifestaient une consommation accrue du feuillage des deux lignées de soja. Les larves parasitées élevées sur les feuilles de la variété résistante GAT “81–327” pesaient moins et produisaient moins également de parasites adultes.

     

Recognition of sex in the acoustic communication of the grasshopper Chorthippus biguttulus (Orthoptera, Acrididae)

Many gomphocerine grasshoppers communicate acoustically: a male's calling song is answered by a female which is approached phonotactically by the male. Signals and recognition mechanisms were investigated in Chorthippus biguttulus with regard to the cues which allow sex discrimination. (1) The stridulatory files on the hindfemur of both sexes are homologous in that they are derived from the same row of bristles, but convergent with respect to the “pegs”. In males the pegs are derived from the bristles, and in females from the wall of the bristle's cup. (2) Male and female songs are generated by similar, probably homologous motor programs, but differ in the duration, intensity, “gappyness” of syllables, risetime of pulses, and the frequency spectra. The hindleg co-ordination during stridulation and the resulting temporal song patterns are less variable in males than in females. (3) For both sexes, recognition of a mate's signal depends on species-specific syllable structure. For males it is essential that the female syllables consist of distinct short pulses, whereas females reject “gappy” syllables. Males strongly prefer “ramped” pulses, females respond to syllables irrespective of steeply or slowly rising ramps. Males react only to the low-frequency component, whereas females prefer spectra containing both, low and high frequency components. Accepted: 20 November 1996     

Correlations between the psychological peculiarities of an individual and the efficacy of a single neurofeedback session (by the EEG characteristics)

Modifications of different EEG rhythms induced by a single neurofeedback session (by the EEG characteristics) directed toward an increase in the ratio of the spectral powers (SPs) of the α vs θ oscillations were compared with the psychological characteristics of the tested subjects (the group included 30 persons). A generally accepted neurofeedback technique was used; the intensity of acoustic white noise served as the feedback signal. EEG potentials were recorded from the C3 and C4 leads. Psychological testing was carried out using Eysenck’s (EPQ), Rusalov’s (OST), and (16 PF) questionnaires. The directions of changes in the SPs of EEG frequency components were found to significantly correlate with some individuality-related peculiarities of the tested subjects. The SP of the δ rhythm correlated with the EPQ scale “neuroticism,” OST scale “social plasticity,” and 16 PF factors H (“parmia”), I (“premsia”), and Q₃ (“self-control of behavior”). The SP of the θ component demonstrated correlations with the OST scales “ergisity,” “plasticity,” and “social temp” and with 16 PF factors M (“autia”), Q₄ (“frustration”), and Q1 (“exvia”). The SP of the α rhythm correlated with 16 PF factors Q₃ (“self-control of behavior”), G (“strength of superEgo”), O (“hypothymia”), L (“protension”), and N (“shrewdness”). The SP of the β rhythm correlated with the OST scale “emotionality,” while that of the γ rhythm showed correlations with the 16 PF indices L (“protension”) and M (“autia”). Changes in the ratio of the α vs θ SPs correlated with the EPQ scale “neuroticism.” Thus, our data confirm the statement that a high individual variability of the results of a single (first in the series) neurofeedback session is to a great extent related to peculiarities of the individual psychological pattern of the subject. Neirofiziologiya/Neurophysiology, Vol. 38, No. 3, pp. 239–247, May–June, 2006.     

Different zooplankton structures in the German Bight

In August 1982, a net of 48 stations with altogether 208 samples was investigated in the eastern German Bight with respect to temperature, salinity, as well as the amount and species composition of the mesozooplankton (>80 μm). The data were arranged into different structures by means of a cluster analysis. Four different clusters were found: (a) a “Wadden sea water” with few holoplankton organisms but a higher amount of spionid larvae; (b) a “German Bight water” with a maximum occurrence of turbellaria (Alaurina composita) and medium concentrations of copepods; (c) a mixing area between these two water masses with highest amounts ofOikopleura dioica, Temora longicornis, Acartia sp., mussel larvae and larvae of the spionid worms; (d) a “North Sea water” mass with highest concentrations ofPseudocalanus elongatus, Paracalanus parvus undOithona similis. The differences in the concentrations of the species mentioned between the four clusters were significant on the 0.1%-level.     

“Camel nanoantibody” is an efficient tool for research,diagnostics and therapy

This short review provides an introduction to the rapidly developing field of generation and utilization of “camel nanoantibodies” (or “nanobodies”). The term “nanoantibody” or “nanobody” was given to single-domain variable fragments of special type of antibodies that naturally exist (in addition to classical types of antibodies) in blood of Camelidae family animals and in some chondrichthyan fishes. The existence of very efficient technology of nanobody generation and some very useful characteristic features promise a big potential for their use in immunobiotechnology and medicine.     

Inheritance of resistance to Plum pox virus in apricot (Prunus armeniaca L.)

The inheritance of resistance to Plum pox virus (PPV) has been studied in 1,178 apricot hybrids. Seven hundred and eighteen F1 hybrids, obtained from controlled crosses between the susceptible Greek cultivar “Bebecou” and the resistant PPV cultivars of American origin (“Stark Early Orange,” ‘NJA2,” ‘Veecot,” “Sunglo,” “Harlayne,” and “Orangered”) were evaluated for resistance to the PPV-M (Marcus) strain, 8 years after artificial inoculation. The inheritance of resistance to PPV has been additionally studied for the first time in a BC₁ population of 95 apricot hybrids for four vegetative periods. Reaction of each hybrid to PPV-M was scored through visual symptoms, indexing onto GF-305 and double-antibody sandwich enzyme-linked immunosorbent assay tests. Segregation within the hybrids, determined by Chi-squared analysis, fits a 1:1 ratio (P ≤ 0.05) of the resistant vs susceptible, indicating that resistance to PPV is controlled by a single dominant gene locus and that the above six resistant cultivars are heterozygous for the trait. Plants carrying this gene may initially develop disease symptoms on leaves but eventually recover and no virus can be detected in leaves. Susceptible plants show similar symptoms initially but remain symptomatic. Inheritance of resistance to PPV also has been studied in 365 F1 hybrids by crossing the resistant cultivar “Stella” with the susceptible “Bebecou” and the resistant cultivars “Sunglo” and “NJA2,” for 8 years after inoculation. The segregation ratio was 1:0 (resistant/susceptible) suggesting that “Stella” is homozygous for the resistance trait. The purpose of this work was the enhancement of the knowledge of inheritance of resistance to PPV for the selection of new cultivars.     

The effects of external potassium and magnesium concentrations on the magnesium and potassium inflow rates and growth of micropropagated cherry rootstocks, “F.12/1” (Prunus avium L.) and “Colt” (Prunus avium L.) × Prunus pseudocerasus L.)

The effects (and interaction) of two solution concentrations of Mg (50, 500, μM) and two of K (250, 4250 μM) on the growth of micropropagated plants of “F. 12/1” and “Colt” were investigated using a flowing solution culture system. Magnesium inflow and growth of “Colt” and “F. 12/1” were inhibited to a similar extent by an increased concentration of K in the nutrient solution. However, the consequences of this inhibition were different. Reduced inflow of Mg in “F. 12/1” caused Mg deficiency symptoms at high and low concentrations of K, whereas this only occurred with a combination of high K concentration and low Mg concentration in “Colt”. The distribution of dry matter within the plant was significant in determining susceptibility to Mg deficiency. Since “F. 12/1” has a smaller root:shoot ratio than Colt it is unable to sustain the same concentration of Mg in leaves as “Colt” irrespective of external K concentration. The molar ratio of K:Mg in soil solutions should remain <8.5:1 in order to ensure maximum growth of “F. 12/1” and “Colt”. This revised version was published online in June 2006 with corrections to the Cover Date.     

Removal of the polar lobe leads to the formation of functionally deficient photocytes in the annelidChaetopterus variopedatus

Summary The light emitting photocytes ofChaetopterus variopedatus larvae are bilaterally situated within the ectoderm of the post-trochal region. Their histological appearance is similar to that of the adult photocytes. The larval photocytes contain a large quantity of membranous secretory vesicles (photosomes), which probably contain the photoluminescent protein. The two-cellChaetopterus embryo contains a small AB and a large CD blastomere. Previous studies have shown that only the “larvae” resulting from isolated CD blastomeres are able to luminesce. Consistent with these findings, morphologically distinct photocytes are only found in the CD larvae. The removal of the small polar lobe that forms during first cleavage leads to the production of a “larva” that is unable to produce light. All delobed larvae contain morphologically distinct photocytes, which are identical to those in normal larvae except they appear to contain only a small quantity of photosomes. Experimental equalization of first cleavage leads to the production of a double embryo. While photocytes are found in both of the duplicated post-trochal regions, usually only one of these is capable of emitting luminescence. Apparently, the highly localized vagetal material (determinants) responsible for functional light emission is distributed to both halves in only a few cases when first cleavage is experimentally “equalized”. These results indicate that the determinative action of the polar lobe is not required for the formation of the photocytes themselves, but rather for their ability to function as emitters of light. The determinants in the polar lobe ofChaetopterus may control some aspect of the photoluminescence reaction itself, such as the production of the photoprotein.     

Exploring Scientific Misconduct: Isolated Individuals,Impure Institutions,or an Inevitable Idiom of Modern Science?

This paper identifies three distinct narratives concerning scientific misconduct: a narrative of “individual impurity” promoted by those wishing to see science self-regulated; a narrative of “institutional impropriety” promoted by those seeking greater external control of science; and a narrative of “structural crisis” among those critiquing the entire process of research itself. The paper begins by assessing contemporary definitions and estimates of scientific misconduct. It emphasizes disagreements over such definitions and estimates as a way to tease out tension and controversy over competing visions of scientific research. It concludes by noting that each narrative suggests a different approach for resolving misconduct, and that the difference inherent in these views may help explain much of the discord concerning unethical behavior in the scientific community.     

Physiological bases of oligotrophy of microorganisms and the concept of microbial community

Three groups of physiological processes in microorganisms are considered the physiological basis of oligotrophy: the greater substrate affinity of the oligotrophs' transport systems, efficient or “economical” metabolism, and existence of a “master reaction” or “rate-determining steps” controlling the rate of metabolism. Heterotrophic microorganisms are divided into three unequal groups according to “reaction norma.” Two groups representing the extremes are small groups with the “narrow” reaction norma, regarding the concentrations and structure of the assimilated organic compounds and variability limits of the physiological characteristics mentioned above. The third, intermediate group includes the majority of microorganisms with the “wide” reaction norma.     

Unreasonable implications of reasonable idiotypic network assumptions

We first analyse a simple symmetric model of the idiotypic network. In the model idiotypic interactions regulate B cell proliferation. Three non-idiotypic processes are incorporated: (1) influx of newborn cells; (2) turnover of cells: (3) antigen. Antigen also regulates proliferation. A model of 2 B cell populations has 3 stable equilibria: one virgin, two immune. The twodimensional system thus remembers antigens, i.e. accounts for immunity. By contrast, if an idiotypic clone proliferates (in response to antigen), its anti-idiotypic partner is unable to control this. Symmetric idiotypic networks thus fail to account for proliferation regulation. In high-D networks we run into two problems. Firstly, if the network accounts for memory, idiotypic activation always propagates very deeply into the network. This is very unrealistic, but is an implication of the “realistic” assumption that it should be easier to activate all cells of a small virgin clone than to maintain the activation of all cells of a large (immune) clone. Secondly, graph theory teaches us that if the (random) network connectance exceeds a threshold level of one interaction per clone, most clones are interconnected. We show that this theory is also applicable to immune networks based on complementary matching idiotypes. The combination of the first “percolation” result with the “interconnectancr” result means that the first stimulation of the network with antigen should eventually affect most of the clones. We think this is unreasonable. Another threshold property of the network connectivity is the existence of a virgin state. A gradual increase in network connectance eliminates the virgin state and thus causes an abrupt change in network behaviour. In contrast to weakly connected systems, highly connected networks display autonomous activity and are unresponsive to external antigens. Similar differences between neonatal and adult networks have been described by experimentalists. The robustness of these results is tested with a network in which idiotypic inactivation of a clone occurs more generally than activation. Such “long-range inhibition” is known to promote pattern formation. However, in our model it fails to reduce the percolation, and additionally, generates semi-chaotic behaviour. In our network, the inhibition of a clone that is inhibiting can alter this clone into a clone that is activating. Hence “long-range inhibition” implies “long-range activation”, and idiotypic activation fails to remain localized. We next complicate this model by incorporating antibody production. Although this “antibody” model statically accounts for the same set of equilibrium points, it dynamically fails to account for state switching (i.e. memory). The switching behaviour is disturbed by the autonomous slow decay of the (long-lived) antibodies. After antigenic triggering the system now performs complex cyclic behaviour. Finally, it is suggested that (idiotypic) formation of antibody complexes can play only a secondary role in the network. In conclusion, our results cast doubt on the functional role of a profound idiotypic network. The network fails to account for proliferation regulation, and if it accounts for memory phenomena, it “explodes” upon the first encounter with antigen due to extensive percolation.     

Pulses of movement across the sea ice: population connectivity and temporal genetic structure in the arctic fox

Lemmings are involved in several important functions in the Arctic ecosystem. The Arctic fox (Vulpes lagopus) can be divided into two discrete ecotypes: “lemming foxes” and “coastal foxes”. Crashes in lemming abundance can result in pulses of “lemming fox” movement across the Arctic sea ice and immigration into coastal habitats in search for food. These pulses can influence the genetic structure of the receiving population. We have tested the impact of immigration on the genetic structure of the “coastal fox” population in Svalbard by recording microsatellite variation in seven loci for 162 Arctic foxes sampled during the summer and winter over a 5-year period. Genetic heterogeneity and temporal genetic shifts, as inferred by STRUCTURE simulations and deviations from Hardy–Weinberg proportions, respectively, were recorded. Maximum likelihood estimates of movement as well as STRUCTURE simulations suggested that both immigration and genetic mixture are higher in Svalbard than in the neighbouring “lemming fox” populations. The STRUCTURE simulations and AMOVA revealed there are differences in genetic composition of the population between summer and winter seasons, indicating that immigrants are not present in the reproductive portion of the Svalbard population. Based on these results, we conclude that Arctic fox population structure varies with time and is influenced by immigration from neighbouring populations. The lemming cycle is likely an important factor shaping Arctic fox movement across sea ice and the subsequent population genetic structure, but is also likely to influence local adaptation to the coastal habitat and the prevalence of diseases.     

An ethological analysis of types of agonistic interaction in a captive group of Java-monkeys (Macaca fascicularis)

The primate literature provides many indications not only that the nature of dyadic interactions is to a large extent determined by the relations of the interacting animals with others and between these others, but also of the existence of polyadic interactions in which more than two individuals are simultaneously involved. The objectives of the present study are to obtain a quantitative categorization of the agonistic interaction types of captive Java-monkeys and an analysis of their dynamics. After having described the agonistic behaviour patterns of Java-monkeys we shall discuss the categorization of agonistic interaction types (depending on the number of involvees: “dyads”, “triads” and “polyads”), the way in which these types can be further differentiated on the basis of the nature and the direction of the behaviours shown (e.g., different types of alliances), and the existence of so-called “sub-directed” behaviours (i.e., non-agonistic behaviours which are shown towards a dominant third animal more or less simultaneously with aggressive behaviour directed towards an opponent). The analysis indicates that agonistic behaviour is different both in its form and its regulation in interactions of different complexity. This research was supported in part by a government grant (i.e.: Beleidsruimte project: 16-21-06, “Brain and Behaviour”) to the first author. The investigation was supported by a grant from the Beleidsruimtemiddelen Hersenen en Gedrag to the first author.     

Peculiarities of Activation of the Shoulder Belt and Shoulder Muscles in Generation of Different-Direction Isometric Efforts by the Forearm

We studied central motor commands, CMCs, coming to the muscles that flex and extend the shoulder and elbow joints in the course of generation of voluntary isometric efforts of different directions by the forearm; the efforts were initiated according to a visual signal. Amplitudes of EMGs recorded from the muscles of the shoulder belt and shoulder and subjected to full-wave rectification and low-frequency filtration were considered correlates of the CMC intensity. An effort of the preset direction was developed within the operational space of the horizontal plane with angles 30 deg in the shoulder joint (external angle with respect to the frontal plane) and 90 deg in the elbow joint. We plotted sector diagrams of the logarithmic coefficient of the intensity increment of EMGs of the above muscles for the entire set of directions of generated efforts with a 15- or 20-deg step. Orientations of the maxima of EMG activity of the given muscles were rather close to the directions of the maxima of the force moments generated by these muscles. In most cases, a shift of the direction by one gradation with respect to the EMG maximum in the respective muscle resulted in a significant decrease in the level of EMG activity. It is shown that preferential activation of the muscles agonistic with respect to the examined direction of the generated effort was, as a rule, accompanied by coactivation of the antagonist muscles. When “two-joint” isometric efforts are formed, realization of the socalled synergic muscle tasks (where prevailing contractions of the muscles of the same functional direction for both joints coincide, i.e., flexion-flexion or extension-extension) is organized in a simpler manner. The programs of “nonsynergic” contractions (flexion of one joint and extension of another one, or vice versa) are more complex. In different subjects, considerably dissimilar patterns of EMG activity in muscles influencing these joints could be observed.     

Chironomidae (Diptera) larvae assemblages differ along an altitudinal gradient and temporal periods in a subtropical montane stream in Northwest Argentina

The objective of this study was to analyze the spatial and temporal dynamics of Chironomidae larvae assemblages in a subtropical mountain river basin of northwestern Argentina. We hypothesized that they would respond to (1) environmental changes along the altitudinal gradient in the Yungas forests; (2) environmental changes caused by spates (e.g., increased discharge or sediment transportation). We sampled five sites along an altitudinal gradient of ca. 1,500 m (from 680 to 2,170 m) during low-water and high-water period for 2 years. The Chironomidae larvae assemblages varied along the altitudinal gradient and between sampling periods based on an indicator species analyses and an ordination analyses (NMDS). The best indicator for high altitude sites was “Genus X” followed by Onconeura sp.2, “Genus 9”, “Genus 10”, and Rheotanytarsus sp.2; while Rheotanytarsus lamellatus, Oliveiriella sanjavieri, and Thienemanniella sp. were good indicators for low altitude sites. O. sanjavieri and R. lamellatus were the best indicators for high-water period. The most relevant environmental variables influencing their spatial and temporal distribution were altitude, water temperature, conductivity, and pH. This study is the first to analyze and describe in detail the spatial and temporal distribution of Chironomidae larvae assemblages in northwestern Argentina, and most of the taxa reported here are new records for the region.     

“Ignition” phenomena in random nets

The spread of excitation in a “random net” is investigated. It is shown that if the thresholds of individual neurons in the net are equal to unity, a positive steady state of excitation will be reached equal to γ, which previously had been computed as the weak connectivity of the net. If, however, the individual thresholds are greater than unity, either no positive steady state exists, or two such states depending on the magnitude of the axone density. In the latter case the smaller of the two steady states is unstable and hence resembles an “ignition point” of the net. If the initial stimulation (assumed instantaneous) exceeds the “ignition point,” the excitation of the net eventually assumes the greater steady state. Possible connections between this model and the phenomenon of the “preset” response are discussed.     

Use of microsatellite DNA and otolith Sr:Ca ratios to infer genetic relationships and migration history of four morphotypes of <Emphasis Type="Italic">Rhinogobius</Emphasis> sp. OR

The genetic diversity and relationship among four morphotypes of Rhinogobius sp. OR, Gobiidae (“Tōshoku,” “Shinjiko,” “Gi-tōshoku,” and “Shimahire”) were investigated with seven microsatellite DNA loci, and amphidromy of these morphotypes was verified by strontium (Sr) and calcium (Ca) deposition in the otolith. Samples of “Tōshoku,” “Shinjiko,” “Gi-tōshoku,” and “Shimahire” were collected from, respectively, three, three, two, and four locations in Japan. Microsatellite analysis detected high genetic diversity (based on the number of alleles, allelic richness, and average observed heterozygosity) in the “Tōshoku” and “Shinjiko” morphotypes relative to the “Shimahire” morphotype; the “Gi-tōshoku” morphotype had an intermediate level of variation. Almost all pairwise F _ST values were significantly different from zero (P < 0.001), except between two populations of “Tōshoku.” Clear genetic independence was observed between the “Shinjiko” and “Shimahire” morphotypes in the Maruyama River. A principle component analysis based on microsatellite data indicated that the “Tōshoku,” “Shinjiko,” and “Gi-tōshoku” morphotypes were genetically similar. Furthermore, the three populations of “Tōshoku” were closely related each other, and two of those collected from the Lake Biwa system were a single population. There was, however, a high degree of genetic differentiation between “Shimahire” and the other morphotypes; moreover, there was high genetic divergence among four populations of the “Shimahire” morphotype. Amphidromous migratory histories were indicated by Sr:Ca ratios in two of three populations of the “Shinjiko” morphotype and in one of two “Gi-tōshoku” morphotypes, whereas all populations of the “Shimahire” morphotype were freshwater residents. The large genetic divergence and low genetic diversity in “Shimahire” are likely related to migration history.