Context and Development of Sexual Behavior of Wild Bonobos (Pan paniscus) at Wamba, Zaire

I studied sexual behavior of immature bonobos (Pan paniscus) in a wild group living at Wamba, Zaire, with special reference to its development. Even immature individuals under 1 year old performed sexual behavior. Sexual behavior occurred in almost all age–sex combinations, except between immature and mature females. Based on analyses of behavioral pattern and context, I classified sexual behavior involving immature individuals into three categories. (1) Genital contact between immature individuals was observed during play, and was performed by males more frequently than by females. This sexual behavior shared many traits with that of other great apes. (2) Copulation-like genital contact was observed between immature males and mature females. Its frequency increased with the immature male's age; it developed into copulation in adulthood. (3) Genital contact used to regulate interindividual relationships. This behavior, which is unique to bonobos, was absent among infants. It developed between late juvenile and early adolescent periods in association with changes in social circumstances.     

Use of a flooded salt marsh habitat by an endangered cyprinodontid fish (<Emphasis Type="Italic">Aphanius iberus</Emphasis>)

We report the first data on the use of occasionally inundated habitats in a Mediterranean coastal lagoon by the Spanish toothcarp (Aphanius iberus), a cyprinodontid fish in danger of extinction. During a flooding period, we sampled the fish population and the macroinvertebrate community in three contrasting habitats: mats of green algae, open water, and a habitat dominated by glasswort (Salicornia patula) that is occasionally inundated. The three habitats displayed strong variation in density and species composition of invertebrates in the water column and the benthos. In general, algal mats had higher invertebrate biomasses, but glasswort had higher diversity of organisms, in part of terrestrial origin. The density of toothcarp was very low in the open water. The habitat that is occasionally inundated (glasswort) significantly had the highest density of mature toothcarp, while immature fish were similarly abundant in the glasswort and algal mat habitats. Condition (weight–length relationship) and total food biomass in the gut contents of immature toothcarp was significantly higher in algal mats than in glasswort, whereas there were no such differences for mature fish. Therefore, the occasionally flooded habitat (glasswort) was positively selected by large mature toothcarp but seemed a disadvantageous habitat for immature individuals. Handling editor: L. Mauricio Bini.     

Reproductive biology and pollen flow of a rheophytic aroid, Furtadoa sumatrensis (Araceae) in the Malesian wet tropics

 A rheophytic aroid, Furtadoa sumatrensis M. Hotta, showed protogynous flowering behaviour, which reduced autogamous pollination within the spadix and promoted outcrossing among spadices by Colocasiomyia sp. (Diptera, Drosophilidae). Self-compatibility of this species allowed geitonogamous crossing between genetically homogeneous spadices from the same individual. Chemical dyes as pollen analogues were carried by Colocasiomyia flies bidirectionally along the forest stream habitat, and their movements were significantly affected by the density of mature spadices in a population. This correlation was consistent among populations. In a population with higher density, the dye's movements were restricted to a smaller distance logarithmically. Estimated neighborhood size (N _e ) and area (A) showed positive and negative correlation with density, respectively. The range of pollen flow was estimated at most within a river system. Received March 7, 2000 Accepted January 23, 2001     

Sexually mature and immature yearling male European ground squirrels: a comparison of behavioral and physiological parameters

Early puberty may increase lifetime reproductive success, but may also entail diverse costs. In male European ground squirrels (Spermophilus citellus), age at sexual maturity varies among individuals. We compared sexually active and inactive yearling males under standardized conditions in outdoor enclosures. Non-reproductive yearling males hibernated significantly longer than mature males and emerged when the mating period had already started. Testosterone concentrations peaked in the precocious males during premating and mating, and were depressed in the non-reproductive individuals throughout the study period. Emergence body mass was similar in both groups, although sexually immature males were significantly heavier at the end of the active season. Additionally, non-reproductive yearlings grew faster than precocious individuals. Male–male aggression peaked during mating and the following 4 weeks. Intense aggressive interactions like chases and fights were mostly initiated by reproductive males and were directed towards mature competitors. Aggressive behavior in immature males was mainly expressed as spatial displacements. Non-agonistic interactions with females were only observed in sexually active males. Concentrations of fecal cortisol metabolites did not differ between the two male groups and were elevated in the second half of the active season. Leucocyte numbers were reduced during mating in mature males, whereas in immature animals, cell numbers did not change significantly. Our results indicate that early puberty in European ground squirrels involves lower growth rates, high intrasexual aggression, lower prehibernatory body mass, and probably worse immunological condition during mating.     

Fecundity and Longevity of Green Vegetable Bug,Nezara viridula, Following Parasitism byTrichopoda giacomellii

The effects of parasitism by the ArgentinianTrichopoda giacomellii(Blanchard) on reproduction and longevity of its host,Nezara viridula(L.) are reported. Parasitoid larvae suppress egg maturation, reducing by 70% the fecundity of mature female hosts during the period of larval development. Egg viability was not affected, but mating frequency was reduced by approximately 50%. When parasitized as newly eclosed adults, 84% of females fail to reproduce. In male hosts, fertility and mating frequency were not affected during the period of larval parasitoid development. In male and reproductively immature female hosts, death was coincident with, or occurred shortly after parasitoid emergence (2–4 days); in mature females, death occurred on average 2 weeks after larval parasitoid emergence. Host mortality occurred as a consequence of tissue damage incurred as the parasitoid larvae emerged from the host. Some individuals survived parasitism though no further reproductive activity (mating or oviposition) occurred. The effectiveness ofT. giacomelliias a biological control agent is discussed in relation to its impact on reproduction and survival of its host and contrasted with the action of otherTrichopodaspecies.     

The reintroduction of the Eurasian otter (Lutra lutra) into the Netherlands: hidden life revealed by noninvasive genetic monitoring

The last recorded presence of the Eurasian otter (Lutra lutra) in the Netherlands dates from 1989 and concerned a dead individual. In 2002 a reintroduction programme was started, and between June 2002 and April 2008 a total of 30 individuals (10 males and 20 females) were released into a lowland peat marsh in the north of the Netherlands. Noninvasive genetic monitoring based on the genetic profiles obtained from DNA extracted from otter faeces (spraints) was chosen for the post-release monitoring of the population. To this end, the founding individuals were genotyped before release and spraints were collected in the release area each winter from 2002 to 2008. From June 2002 to April 2008 we analysed the genetic profile of 1,265 spraints on the basis of 7–15 microsatellite loci, 582 of which (46%) were successfully assigned to either released or newly identified genotypes. We identified 54 offspring (23 females and 31 males): the females started to reproduce after 2 years and the males after 4 years. The mating and reproductive success among males was strongly skewed, with a few dominant males fathering two-thirds of the offspring, but the females had a more even distribution. The effective population size (Ne) was only about 30% of the observed density (N), mainly because of the large variance in reproductive success among males. Most juvenile males dispersed to surrounding areas on maturity, whereas juvenile females stayed inside the area next to the mother’s territory. The main cause of mortality was traffic accidents. Males had a higher mortality rate (22 out of 41 males (54%) vs. 9 out of 43 females (21%)). During winter 2007/08 we identified 47 individuals, 41 of which originated from mating within the release area. This study demonstrates that noninvasive molecular methods can be used efficiently in post-release monitoring studies of elusive species to reveal a comprehensive picture of the state of the population.     

Evolution of mating systems in coral reef gobies and constraints on mating system plasticity

Social and mating systems can be influenced by the distribution, abundance, and economic defendability of breeding partners and essential resources. Polygyny is predicted where males can economically defend multiple females or essential resources used by females. In contrast, monogamy is predicted where neither sex can monopolise multiple partners, either directly or through resource control, but where one mate is economically defendable. The mating system and reproductive behaviour of five species of coral reef goby were investigated and contrasted with population density and individual mobility. The two most abundant species (Asterropteryx semipunctatus and Istigobius goldmanni) were polygynous. In contrast, the less populous and more widely dispersed epibenthic species (Amblygobius bynoensis, Amblygobius phalaena and Valenciennea muralis) were pair forming and monogamous. All five species had low mobility, mostly remaining within metres (3 epibenthic species) or centimetres (2 cryptobenthic species) of a permanent shelter site. Interspecific differences in the mating system may have been shaped by differences in population density and the ability of reproductive individuals to economically defend breeding partners/sites. However, in a test of mating system plasticity, males of the three monogamous species did not mate polygynously when given the opportunity to do so in experimental manipulations of density and sex ratio. Mate guarding and complex spawning characteristics, which have likely co-evolved with the monogamous mating system, could contribute to mating system inflexibility by making polygynous mating unprofitable for individuals of the pair forming species, even when presented with current-day ecological conditions that usually favour polygyny.     

Genotypic and phenotypic analyses reveal distinct population structures and ecotypes for sugar beet‐associated Pseudomonas in Oxford and Auckland

Fluorescent pseudomonads represent one of the largest groups of bacteria inhabiting the surfaces of plants, but their genetic composition in planta is poorly understood. Here, we examined the population structure and diversity of fluorescent pseudomonads isolated from sugar beet grown at two geographic locations (Oxford, United Kingdom and Auckland, New Zealand). To seek evidence for niche adaptation, bacteria were sampled from three types of leaves (immature, mature, and senescent) and then characterized using a combination of genotypic and phenotypic analysis. We first performed multilocus sequence analysis (MLSA) of three housekeeping genes (gapA, gltA, and acnB) in a total of 152 isolates (96 from Oxford, 56 from Auckland). The concatenated sequences were grouped into 81 sequence types and 22 distinct operational taxonomic units (OTUs). Significant levels of recombination were detected, particularly for the Oxford isolates (rate of recombination to mutation (r/m) = 5.23 for the whole population). Subsequent ancestral analysis performed in STRUCTURE found evidence of six ancestral populations, and their distributions significantly differed between Oxford and Auckland. Next, their ability to grow on 95 carbon sources was assessed using the Biolog? GN2 microtiter plates. A distance matrix was generated from the raw growth data (A₆₆₀) and subjected to multidimensional scaling (MDS) analysis. There was a significant correlation between substrate utilization profiles and MLSA genotypes. Both phenotypic and genotypic analyses indicated presence of a geographic structure for strains from Oxford and Auckland. Significant differences were also detected for MLSA genotypes between strains isolated from immature versus mature/senescent leaves. The fluorescent pseudomonads thus showed an ecotypic population structure, suggestive of adaptation to both geographic conditions and local plant niches.     

Density effects of flowering phenology and mating potential in Nicotiana alata

Summary We investigated effects of plant density on floral phenology and potential mating in artificial populations of the outcrossing ornamental Nicotiana alata planted at three densities. Path analysis revealed that increasing plant density yielded significantly earlier peak flowering dates, significantly earlier last flowering dates, and significantly lower plant biomass. Direct effects of density on final flower number were not significant. Variation among replicate plots for first date of flowering was larger than variation among densities, indicating that factors other than density influence floral initiation.We did not record actual mating, but determined from phenological data the number and identity of potential mates. Increased density had several effects on potential mating patterns and on potential N_e, effective population number. At high density, fewer focal plants flowered for shorter durations. This led to less overlap in flowering time among plants, decreasing the number of potential parental combinations possible among the progeny. Two outcomes of high density, the lower total number of plants flowering and the lower number of plants flowering at most census dates, tended to reduce potential N_e. In contrast, it was low density, where variance in flower number was greatest, that was most likely to yield the greatest reduction in N_e due to variance in progeny number.At high density the potential for assortative mating among tall plants was much greater and occurred later than among large plants at low density. Much of the potential high density assortative mating occurred late in the phenology of individual plants, when there was likely to be lower fruit set.We discuss how ecological agents that alter flowering phenology can potentially alter the genetics of populations, the level and timing of assortative mating and, if genetic variation for response to such ecological agents exists, the potential selection regime.     

High resolution analysis of mating systems: inbreeding in natural populations of Pinus radiata

Pinus radiata has a history of population bottlenecks and is currently restricted to five relatively small populations, three in mainland California, and two on islands off the coast of Baja California. Using highly polymorphic microsatellite markers and a newly developed statistical approach, we were able to estimate individual inbreeding coefficients and can thus analyse the mating system with high resolution. We find a bimodal distribution of inbreeding coefficients: most individuals result from selfing whereas few (in the mainland populations) to a modest number (in the island populations) are likely selfed. In most other pine species and presumably in the ancestral P. radiata population, occurrence of mature selfed individuals would be impossible because of the high genetic load. We therefore conclude that inbreeding depression has been purged in P. radiata and that the mating system has changed as a consequence.     

Maturation of female common snook Centropomus undecimalis: implications for managing protandrous fishes

The assumption for hermaphroditic fish species that mature individuals of the terminal sex arise directly from mature individuals of the primary sex has led to the use of sex ratios as a proxy for age at maturity (A₅₀). The timing of transition and deficient energy reserves, however, can result in a delay between transition and spawning. To test the assumption of female maturity and investigate the relationship between maturation and energy reserves, common snook, Centropomus undecimalis, a protandrous hermaphrodite, were collected from rivers, estuaries, inlets and offshore habitats on the east coast of Florida during 2010–2015. Immature females were observed every month, with lowest proportions during the peak spawning months of July and August. When calculated based on sex ratio, A₅₀ (8.1 years) overestimated the age at which 50% of the female population was, in fact, mature (4.1–4.9 years). Best-fit models indicate that mesenteric fat index (I_F) and hepato-somatic index (I_H) were significantly affected by gonad phase, month and size and weakly by habitat. In post hoc analysis, immature female I_F did not differ significantly from developing and regenerating females, but immature female I_H was significantly lower than that for all mature phases except animals in the regressing phase. Although immature females may have sufficient energy in terms of fat, it appears that energy is not allocated to reproductive processes, as evidenced by lower I_H. Nonetheless, approximately 95% of females were spawning-capable during peak spawning months, suggesting that the energy threshold at which immature females reach maturity is met by most females each spawning cycle.     

Differences in Mating Propensity Between Immature Female Color Morphs in the Damselfly Ischnura elegans (Insecta: Odonata)

Female-limited color polymorphisms occur in a variety of animal taxa where excessive male sexual harassment may explain the coexistence of multiple female color morphs. In the color polymorphic damselfly Ischnura elegans, mature and immature female color morphs coexist at the mating site where males are in search for suitable mating partners. Here, we study male preference and female mating propensity for the two immature female morphs. As would be expected, compared to mature morphs, both immature female morphs mate much less. Within immature females, one morph consistently mates more frequently compared to the other morph, a pattern that is similar for the ontogenetically corresponding mature female morphs. Preference experiments with the two differently colored immature female morphs, however, did not indicate male mate preference for either morph. Low mating frequencies of immature females at natural sites in combination with relatively high attractiveness of immature models in terms of male preference indicate that female behavior influences female mating success.     

Preliminary insights into the spatial ecology and movement patterns of a regionally critically endangered skate (Rostroraja alba) associated with a marine protected area

ABSTRACT

The implementation of spatial protection measures is currently a priority in batoid species' conservation strategies, but their spatial ecology remains largely unknown. We provide some preliminary insights into the movement patterns of the white skate (Rostroraja alba), a batoid classified as Critically Endangered in European waters. Three individuals (two females: one mature, one immature; and one immature male) were monitored with acoustic telemetry in a marine protected area (Portugal). The mature female remained present in the study area throughout the whole monitoring period (20 months). Residency analyses revealed this specimen spent more than 70% of the time within this coastal marine park. The immature female and the immature male were only detected during three and four months, respectively. Whether the uncovered movement patterns are common within the population remains unclear, but the present study provides useful information to better plan future research on the movement ecology of a rather unstudied species.     

Intraspecific resource partitioning by an opportunistic strategist, inland silverside Menidia beryllina

Seasonal variation in prey consumption and food resource overlap was evident in an inland water body for mature male, mature female and immature inland silverside (Menidia beryllina). During the first growth phase marked by intensive somatic growth by immature inland silverside, few adults were present in the population (28% of total catch), thus minimizing intraspecific competition for food resources between juvenile and adult inland silverside. During the second growth phase by adults, few juvenile inland silverside were present (0% of total catch) in the population, again minimizing intraspecific competition for food resources between juvenile and adult inland silverside. A divergence in food resource overlap was observed when mature male, mature female and immature inland silverside were present in the population. These population‐level demographic responses to energy acquisition are likely necessary to maximize individual growth of mature male, mature female and immature inland silverside.     

Sex ratio and maturity indicate the local dispersal and mortality of adult stoneflies

1. Despite a recent upsurge in interest, there remains remarkably little information about the dispersal and survival of the adults of most stream‐dwelling insects, although this is a basic requirement for understanding their long‐term population dynamics. 2. Using Malaise traps for a whole annual flight period, we captured adult stoneflies (Leuctra nigra) along transects perpendicular to three upland Welsh streams. We assessed spatial and temporal patterns in sex ratio to infer local dispersal and, using maturity as an age marker, estimated the mortality of adult females. 3. Nearly all adult stoneflies (99%) were taken in the period 26 April–23 July, and the onset of the male and female flight periods was the same. Most males (90%) had been caught by late June. Females were classified as immature (without ripe eggs) or mature, and 90% of immature females had been caught by mid‐late June (depending on catchment). As immature females declined in the catch, mature females increased, 10% having been caught by late‐May to early June and 90% by early to mid‐July. The median catches of immature and mature females were separated by 32 days in all three catchments. 4. There was a female bias in the sex ratio overall, which increased as time passed and was attributable partially to the greater longevity of females. Late in the flight period, however, female bias was also greater near the stream channel implying a return of mature females (but not males) from the riparian vegetation, presumably to oviposit. 5. The number of mature females was less than the number of immatures in two of the three channels. Over all three catchments, the overall mortality of females over the 32 days taken to mature was estimated at 29%, equivalent to an exponential daily rate of 1.1% day^?1. The apparently negative mortality rate in one catchment (i.e. more matures than immatures being caught) could be due to an influx of adult females from elsewhere along the channel to oviposit. 6. Natural markers of age and population structure, such as sex ratio and female maturity, thus enabled us to detect a return of females to the stream to oviposit, after prior limited dispersal into the riparian zone, and to infer longitudinal movements in search of suitable sites. We were also able to estimate mortality in the field. Such natural markers seem to have been underexploited in the study of adult aquatic insects.     

Isolation and characterization of novel tetranucleotide microsatellite DNA markers for the spotted salamander,Ambystoma maculatum

Fifteen tetranucleotide microsatellite loci were identified and characterized for spotted salamanders (Ambystoma maculatum) collected from three vernal pools in the south‐eastern USA. These markers revealed a high degree of genetic diversity (7–32 alleles per locus), heterozygosity (31.6–86.3%) and allelic heterogeneity (91% of comparisons were statistically significant). Considerable differentiation among populations was observed as genetic distances (chord) ranged between 0.50 and 0.65, and all F_ST values (0.08–0.14) were statistically significant. Moreover, genotypic assignment tests correctly classified all individuals to their respective collection. These markers should prove useful for investigating fine‐scale population structure and mating system.     

Evolution of mating types in finite populations: The precarious advantage of being rare

Sexually reproducing populations with self‐incompatibility bear the cost of limiting potential mates to individuals of a different type. Rare mating types escape this cost since they are unlikely to encounter incompatible partners, leading to the deterministic prediction of continuous invasion by new mutants and an ever‐increasing number of types. However, rare types are also at an increased risk of being lost by random drift. Calculating the number of mating types that a population can maintain requires consideration of both the deterministic advantages and the stochastic risks. By comparing the relative importance of selection and drift, we show that a population of size N can maintain a maximum of approximately N^1/3 mating types for intermediate population sizes, whereas for large N, we derive a formal estimate. Although the number of mating types in a population is quite stable, the rare‐type advantage promotes turnover of types. We derive explicit formulas for both the invasion and turnover probabilities in finite populations.     

Sexually mature and immature yearling male European ground squirrels: A comparison of behavioral and physiological parameters

Early puberty may increase lifetime reproductive success, but may also entail diverse costs. In male European ground squirrels (Spermophilus citellus), age at sexual maturity varies among individuals. We compared sexually active and inactive yearling males under standardized conditions in outdoor enclosures. Non-reproductive yearling males hibernated significantly longer than mature males and emerged when the mating period had already started. Testosterone concentrations peaked in the precocious males during premating and mating, and were depressed in the non-reproductive individuals throughout the study period. Emergence body mass was similar in both groups, although sexually immature males were significantly heavier at the end of the active season. Additionally, non-reproductive yearlings grew faster than precocious individuals. Male–male aggression peaked during mating and the following 4 weeks. Intense aggressive interactions like chases and fights were mostly initiated by reproductive males and were directed towards mature competitors. Aggressive behavior in immature males was mainly expressed as spatial displacements. Non-agonistic interactions with females were only observed in sexually active males. Concentrations of fecal cortisol metabolites did not differ between the two male groups and were elevated in the second half of the active season. Leucocyte numbers were reduced during mating in mature males, whereas in immature animals, cell numbers did not change significantly. Our results indicate that early puberty in European ground squirrels involves lower growth rates, high intrasexual aggression, lower prehibernatory body mass, and probably worse immunological condition during mating.     

Fighting Tactics of Fallow Bucks (Dama dama,Cervidae): Reducing the Risks of Serious Conflict

Competition between male fallow deer (Dama dama), during the breeding season was studied to determine if conflict strategies were consistent with the reduction of risk. Agonistic interactions between males were analysed in relation to age, dominance rank and availability of mating opportunities. The breeding season was divided into two main periods: the pre-rut began when all males had cleaned the velvet from their antlers and ended on the last day before matings were first observed, while the rut refers to the period between the first and last matings. Overall, socially mature males (≥4yr old) were involved in more interactions than immature (≤3yr old) males. Males established dominance rank largely by non-contact agonistic interactions during the pre-rut and there was substantial carry-over of rank to the rut, when it was correlated with mating success. The mating success of males was skewed; mature males achieved 99.4% of the matings and immature males accounted for 0.6%. A mature male was 13 times more likely to fight than an immature male; the mature males that fought most often did so between 0.4 and 0.5 times per hour. During the rut, the number of fights was positively correlated across days with the number of matings. The majority of agonistic interactions (79%) comprising dyads of immature males, involved antler contact. In contrast, mature males engaged antlers in only 42% of their interactions. Fights between mature males lasted more than twice as long as those between immature males and were more likely to occur between opponents with similar dominance ranks. However, towards the end of the rut formerly mismatched opponents were more likely to fight. Thus males operated conditional competitive strategies to decide when to interact and fight. The persistence of rank order from the pre-rut period to the rut and the tendency for mature males to resolve disputes without antler contact, served to reduce the frequency of fights and therefore the risk of serious injury.     

Assortative mating and differential male mating success in an ash hybrid zone population

Background  

The structure and evolution of hybrid zones depend mainly on the relative importance of dispersal and local adaptation, and on the strength of assortative mating. Here, we study the influence of dispersal, temporal isolation, variability in phenotypic traits and parasite attacks on the male mating success of two parental species and hybrids by real-time pollen flow analysis. We focus on a hybrid zone population between the two closely related ash species Fraxinus excelsior L. (common ash) and F. angustifolia Vahl (narrow-leaved ash), which is composed of individuals of the two species and several hybrid types. This population is structured by flowering time: the F. excelsior individuals flower later than the F. angustifolia individuals, and the hybrid types flower in-between. Hybrids are scattered throughout the population, suggesting favorable conditions for their local adaptation. We estimate jointly the best-fitting dispersal kernel, the differences in male fecundity due to variation in phenotypic traits and level of parasite attack, and the strength of assortative mating due to differences in flowering phenology. In addition, we assess the effect of accounting for genotyping error on these estimations.