Dorsal skin color patterns among southern right whales (Eubalaena australis): genetic basis and evolutionary significance.

Distribution and inheritance of dorsal skin color markings among two populations of southern right whales (Eubalaena australis) suggest that two genes influence dorsal skin color. The grey-morph and partial-grey-morph phenotypes (previously known as partial albino and grey-blaze, respectively) appear to be controlled by an X-linked gene, whereas the white blaze appears controlled by an autosomal gene (recessive phenotype). Calving intervals, calf size, and length of sighting history data suggest that partial-grey-morph, white-blaze, and black cows experience similar levels of reproductive success. Grey-morph cows (XgXg) are rare or absent in the two populations, but this was not unexpected given observed population frequencies of grey-morph males (XgY) and partial-grey-morph females (XGXg). The proportion of partial-grey-morph calves produced by black cows (XGXG) suggests that the reproductive success of grey-morph males was equal to that of black males, however, larger sample sizes are required to determine whether grey-morph males tend to have shorter sighting histories. The reproductive success of white-blaze males appeared similar to that of black males among whales off Argentina. There were significantly fewer white-blaze calves than expected off South Africa, which could be due to white-blaze males experiencing reduced reproductive success or to sighting blases that result in white-marked calves being misidentified as black calves. The relative frequencies of both types of dorsal color markings varied between the South African and Argentinian right whale populations, suggesting limited nuclear gene flow between these populations; analyses using other nuclear markers are under way to confirm the extent of gene flow.     

A small badge of longevity: opposing survival selection on the size of white and black wing markings

According to handicap principle, exaggerated ornamental traits are supposed to exert costs on their bearers. However, there is much less theoretical and practical consensus about whether and under which conditions ornament expression should positively correlate with survival. We measured age‐related variation and survival selection on the size of white wing patches and black wing tips in a long‐lived monogamous seabird, the common gull Larus canus. Males had larger white patches than females but patch size showed concave relationship with age irrespective of sex, suggesting that white patch size was prone to senescence in both sexes. Extent of wing tip abrasion correlated negatively with the size of white patch, suggesting, in agreement with the Zahavian handicap hypothesis that only individuals with largest ornaments are able of maintaining them and not paying cost of displaying them. Areas of white wing patches and black wing tips correlated negatively. Irrespective of sex, survival selection favored birds with larger white wing patches and smaller black wing tips, which suggests that white and black wing markings may have coevolved as reverse components of a single ornament. Altogether, our results provide an evidence for the case where survival selection on ornamental traits in females is not weaker than in males. Absence of sex differences with respect to most of observed patterns is consistent with a prediction that among monogamous long‐lived species with biparental care, mutual mate choice leads to evolution of elaborate ornamental traits in both sexes.     

The production of chimeric rats and their use in the analysis of the hooded pigmentation pattern

New, improved media and procedures for making rat chimeric embryos and culturing them in vitro have been developed. We have produced 27 rat chimeras: 20 males and 7 females. This ratio of males to females is consistent with that seen in mouse chimeras, suggesting that rat sex chimeras develop as phenotypic males. By aggregating embryos containing appropriate genetic markers for pigment cell differentiation, it is possible to produce chimeras that elucidate the site of action of the hooded gene. The coat color patterns of black ? black hooded chimeras display a white belly spot. In black ? albino hooded chimeras, small patches of white hair appear on the head and a large white spot occurs on the belly. Black ? agouti hooded chimeras display both agouti and nonagouti pigmentation over the entire surface of the chimera. These animals are fully pigmented with no white spots. In black ? albino non-hooded chimeras, rather small irregular patches of black and white hairs are distributed throughout the pelage. Histological examination of sections of hair follicles obtained from the white areas in the head of black ? albino hooded chimeras revealed amelanotic melanocytes. On the other hand, hair bulbs from the white belly spots do not contain any such melanocytes. Thus the white hairs of the head are due to the presence of albino melanocytes, but the white hairs of the belly are due to the total absence of melanocytes. All these observations are consistent with the conclusion that the hooded gene acts within melanoblasts, probably to retard their migration from the neural crest and/or to prevent their entrance into the hair follicles of the white areas of hooded rats.     

Estimated calf and perinatal mortality in western North Atlantic right whales (Eubalaena glacialis)

Reduced reproductive success has contributed to lack of recovery of the endangered western North Atlantic right whale (Eubalaena glacialis). Here we examined the specific life history period from just before birth through the first year to estimate calf and perinatal losses between 1989 and 2003. The lower bound estimate (17 mortalities from 208 calving events) included documented calf mortalities and presumed deaths from serious injury or disappearance from the sighting record. The upper bound estimated potential calf losses from females with delayed first parturition (>10 yr) and shortened (2 yr) or lengthened (≥4 yr) calving intervals, if the female migrated to the calving ground during these intervals. Because cows were sighted in the calving ground predominantly in years when they were available to calve, adult females sighted there in a possible calving year without a calf were assumed to have experienced a perinatal loss. Twenty-eight potential perinatal losses were detected, bringing the upper bound of calf and perinatal mortality to 45 (3.0 calves/yr). The high frequency of lengthened calving intervals in E. glacialis suggests that abortion and neonatal losses are contributing to lower reproductive success compared to Southern Hemisphere right whales (Eubalaena australis).     

Social monogamy vs. polyandry: ecological factors associated with sex roles in two closely related birds within the same habitat

Why mainly males compete and females take a larger share in parental care remains an exciting question in evolutionary biology. Role‐reversed species are of particular interest, because such ‘exceptions’ help to test the rule. Using mating systems theory as a framework, we compared the reproductive ecology of the two most contrasting coucals with regard to sexual dimorphism and parental care: the black coucal with male‐only care and the biparental white‐browed coucal. Both species occur in the same lush habitat and face similar ecological conditions, but drastically differ in mating system and sexual dimorphism. Black coucals were migratory and occurred at high breeding densities. With females being obligatory polyandrous and almost twice as heavy as males, black coucals belong to the most extreme vertebrates with reversed sexual dimorphism. Higher variance in reproductive success in fiercely competing females suggests that sexual selection is stronger in females than in males. In contrast, resident white‐browed coucals bred at low densities and invariably in pairs. They were almost monomorphic and the variance in reproductive success was similar between the sexes. Black coucals were more likely to lose nests than white‐browed coucals, probably facilitating female emancipation of parental care in black coucals. We propose that a combination of high food abundance, high population density, high degree of nest loss and male bias in the adult sex ratio represent ecological conditions that facilitate role reversal and polyandry in coucals and terrestrial vertebrates in general.     

Colour variation and alternative reproductive strategies in females of the common lizard Lacerta vivipara

Within-sex colour variation is a widespread phenomenon in animals that often plays a role in social selection. In males, colour variation is typically associated with the existence of alternative reproductive strategies. Despite ecological conditions theoretically favourable to the emergence of such alternative strategies in females, the social significance of colour variation in females has less commonly been addressed, relative to the attention given to male strategies. In a population of the common lizard, females display three classes of ventral colouration: pale yellow, orange and mixed. These ventral colours are stable through individual's life and maternally heritable. Females of different ventral colourations displayed different responses of clutch size, clutch hatching success and clutch sex-ratio to several individual and environmental parameters. Such reaction patterns might reflect alternative reproductive strategies in females. Spatial heterogeneity and presence of density- and frequency-dependent feedbacks in the environment could allow for the emergence of such alternative strategies in this population and the maintenance of colour variation in females.     

Factors shaping the evolution of colour patterns in Australian agamid lizards (Agamidae): a comparative study

Identifying general patterns of adaptive coloration in animals can help to elucidate the evolutionary processes that generate them. We examined the evolution of colour patterns in Australian agamid lizards, a morphologically and ecologically diverse group that relies primarily on visual communication. We tested whether certain types of colour (yellow–reds and black) were likely to be used as sexual signals, as indicated by their association with indices of sexual selection, namely, sexual dichromatism and sexual dimorphism in body size and head shape. We then tested whether sexually dichromatic colours are associated with specific patterns (uniform, mottled, striped, blotched, reticulated) or ecological variables such as habitat openness, arboreality, and substrate type. The presence of yellow–red on lateral and ventral body regions and black on ventral body regions was significantly more common in males than females. Lateral yellow–red in males was associated with the total extent of sexual dichromatism and size dimorphism, whereas ventral yellow–red was associated with sexual dichromatism. Both lateral and ventral yellow–red were associated with uniform patterning, suggesting that sexual signals in male agamid lizards may often comprise uniform patches or flushes of yellow–red. Although ventral black coloration was more prevalent on males (i.e. strongly sexually dichromatic), it was not associated with indices of sexual selection, suggesting that, in agamid lizards, yellow–red coloration is more likely to be sexually selected than black. Sexually dichromatic coloration was not strongly associated with any of the ecological variables measured. We found some associations, however, between female dorsal patterns and ecological variables, suggesting that female patterns are influenced by natural selection. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 101–112.     

Genetic evidence for spatio‐temporal changes in the dispersal patterns of two sympatric African colobine monkeys

Western black‐and‐white colobus and Temmink's red colobus are two forest‐dependent African primates with similar ecological requirements, often found in sympatry. Their most striking difference lies in their social system: black‐and‐white colobus live in small groups with mainly male‐mediated dispersal but where females can also disperse, whereas red colobus live in larger groups with males described as philopatric. To investigate whether genetic evidence supports the reported patterns of dispersal based on observational data, we examined eight black‐and‐white and six red colobus social groups from Cantanhez National Park, Guinea‐Bissau. Microsatellite markers revealed a lack of sex‐biased dispersal for black‐and‐white colobus. Gene flow, mainly mediated by females, better explained the genetic patterns found in red colobus, with some evidence for less extensive male dispersal. In contrast to the microsatellite data, low mitochondrial diversity for the black‐and‐white colobus suggests that historical and/or long‐range male‐mediated gene flow might have been favored. In red colobus, the co‐existence of three divergent mitochondrial haplogroups suggests that the Cantanhez population contains a secondary contact zone between divergent lineages that evolved in allopatry. Female‐biased dispersal in this species may be a major factor contributing to the colonization by such differentiated mitochondrial lineages in the region. Overall, we find evidence for a spatio‐temporal change in the dispersal patterns of the colobus monkeys of Cantanhez, with mitochondrial DNA indicating dispersal by mainly a single sex and microsatellite data suggesting that recently both sexes appear to be dispersing within the population. Am J Phys Anthropol, 2013. © 2013 Wiley Periodicals, Inc.     

Reproduction of the Atlantic bobtail squid Sepiola atlantica (Cephalopoda: Sepiolidae) in northwest Spain

Atlantic bobtail squid (Sepiola atlantica) in northwest Spain show seasonal variation in population structure, with juvenile abundance peaking during summer and autumn. However, whether similar patterns exist for reproduction is unknown. Therefore, we describe the reproductive biology of 505 specimens of S. atlantica collected monthly during two consecutive years at two different sites off of Areamilla beach in the Ría de Vigo. Mature males displayed a type of sexual dimorphism previously unknown in members of this species, developing a muscular nodule in the base of each of the ventral arms over ontogeny. Reproductive output of both sexes was similar to that of other bobtail squids. Relative oocyte size (~10% mantle length) appeared to be similar to those of other bobtail squids. Females did not show evidence of having mated before complete maturity. Females of S. atlantica have group‐synchronous ovary maturation, with a positive correlation between female mantle length and ripe oocyte mass, suggesting a terminal investment strategy. Atlantic bobtail squids displayed the same seasonal patterns of reproductive traits at both sampling sites, with significant differences in reproductive activity between males and females. We consider reproductive traits in these small animals as adaptations to the coastal shelf lifestyle.     

Life History Trade-Offs and Behavioral Sensitivity to Testosterone: An Experimental Test When Female Aggression and Maternal Care Co-Occur

Research on male animals suggests that the hormone testosterone plays a central role in mediating the trade-off between mating effort and parental effort. However, the direct links between testosterone, intrasexual aggression and parental care are remarkably mixed across species. Previous attempts to reconcile these patterns suggest that selection favors behavioral insensitivity to testosterone when paternal care is essential to reproductive success and when breeding seasons are especially short. Females also secrete testosterone, though the degree to which similar testosterone-mediated trade-offs occur in females is much less clear. Here, I ask whether testosterone mediates trade-offs between aggression and incubation in females, and whether patterns of female sensitivity to testosterone relate to female life history, as is often the case in males. I experimentally elevated testosterone in free-living, incubating female tree swallows (Tachycineta bicolor), a songbird with a short breeding season during which female incubation and intrasexual aggression are both essential to female reproductive success. Testosterone-treated females showed significantly elevated aggression, reduced incubation temperatures, and reduced hatching success, relative to controls. Thus, prolonged testosterone elevation during incubation was detrimental to reproductive success, but females nonetheless showed behavioral sensitivity to testosterone. These findings suggest that the relative importance of both mating effort and parental effort may be central to understanding patterns of behavioral sensitivity in both sexes.     

Group size and composition influence male and female reproductive success in black howler monkeys (Alouatta pigra)

It has been argued that grouping patterns might influence the reproductive performance of individuals. Increasing group size results in greater travel costs and competition over depletable food resources, which could lead to reduced individual reproductive success. However, in groups with an increasing number of males, female reproductive success is predicted to augment because larger male groups might better protect immatures from infanticidal attacks. In contrast, male reproductive success is predicted to decrease with number of males in a group because fertilization cannot be shared between males. In this paper, we test these predictions on the Mesoamerican black howler monkey (Alouatta pigra) with data on group size and composition for 120 groups from eight populations of black howler monkeys existing in eight protected forests in Mexico and Guatemala. Male and female reproductive success were calculated as a deviation of the observed number of infants (or immatures) from the expected number of infants (or immatures), relative to the number of males and females in a group. Results indicate that both male and female reproductive success decreased with group size. Male reproductive success decreased with an increasing number of males in a group and with increasing proportion of males relative to females in a group. Decreased female reproductive success was associated with increasing number of females in a group, and female reproductive success had a tendency to increase with increasing number of males in a group. These results suggest that in black howler monkeys, living in larger groups might negatively affect the reproductive success of each member. Our findings are similar to those reported for a population of a sister species, Alouatta palliata, living in larger groups.     

Unimodal Batesian polymorphism in the Neotropical swallowtail butterfly Eurytides lysithous (Hbn.)

The Brazilian swallowtail Eurytides lysithous is evidently a Batesian mimic of several Parides species. It is polymorphic for mimetic patterns in both sexes. Various populations contain from one to three major forms, and these seem to depend on two unlinked loci or supergenes. Samples from natural populations, and one reared brood, suggest that one locus controls two white-marked forms, with incomplete dominance producing a third heterozygous form. The heterozygotes are everywhere deficient from Hardy-Weinberg expectations. The second putative locus has an allele epistatic to the first locus which converts the white-marked forms to black, but epistasis is apparently incomplete in heterozygotes. The incomplete dominance and epistasis result in extraordinarily variable polymorphic populations and would allow a genetic analysis for comparison with those already done in the classic Batesian polymorphic swallowtails of the Old World.     

Multitrait aposematic signal in Batesian mimicry

Batesian mimics can parasitize Müllerian mimicry rings mimicking the warning color signal. The evolutionary success of Batesian mimics can increase adding complexity to the signal by behavioral and locomotor mimicry. We investigated three fundamental morphological and locomotor traits in a Neotropical mimicry ring based on Ithomiini butterflies and parasitized by Polythoridae damselflies: wing color, wing shape, and flight style. The study species have wings with a subapical white patch, considered the aposematic signal, and a more apical black patch. The main predators are VS‐birds, visually more sensitive to violet than to ultraviolet wavelengths (UVS‐birds). The white patches, compared to the black patches, were closer in the bird color space, with higher overlap for VS‐birds than for UVS‐birds. Using a discriminability index for bird vision, the white patches were more similar between the mimics and the model than the black patches. The wing shape of the mimics was closer to the model in the morphospace, compared to other outgroup damselflies. The wing‐beat frequency was similar among mimics and the model, and different from another outgroup damselfly. Multitrait aposematic signals involving morphology and locomotion may favor the evolution of mimicry rings and the success of Batesian mimics by improving signal effectiveness toward predators.     

Mating system and reproductive skew in the black rhinoceros

Only approximately 2600 black rhinoceros survive today, mainly in small, isolated populations of < 100 animals. The management of remaining black rhinoceros populations aims at preserving natural levels of genetic relatedness and optimizing breeding success, which requires an accurate knowledge of the mating system, reproductive skew and effective population size. DNA was extracted from faecal samples from a community of 35 wild black rhinoceros, and microsatellites were used to characterize patterns of paternity of 19 offspring born from eight females in this community. Paternity could be ascribed unequivocally for each offspring. Although our conclusions must be considered tentative, we present the first genetic evidence that black rhinoceros males are polygynous, with a high variance in reproductive success. We also describe a noninvasive management tool that can be used for the genetic management of this critically endangered species, both in the wild and in captivity.     

Variable reproductive success in fragmented populations

Marine habitats are naturally patchy and anthropogenic disturbance can further fragment them. Many marine animals are sessile as adults or obligate inhabitants of particular habitats, so populations living in isolated patches of habitat are linked largely by dispersal of planktonic larvae. Theoretically, larvae are more likely to find and settle into large patches of habitat than small patches, thus small habitat patches may experience a more discontinuous supply of recruits resulting in small populations with unusual size- or age-structures or odd sex ratios — conditions where Allee effects on reproductive success are likely. We tested this hypothesis for the Caribbean spotted spiny lobster (Panulirus guttatus), an obligate inhabitant of coral patch reefs whose mating dynamics are size-dependent. We found that P. guttatus were less abundant on small reefs where their size structure and per capita reproductive success were significantly more variable, particularly among large females that are susceptible to sperm limitation that diminishes fertilization rates. These results are indicative of Allee effects and provide a mechanistic understanding of how size-dependent mating dynamics influence reproductive success in ways that alter population dynamics in patchy habitats.     

Female survival,lifetime reproductive success and mating status in a passerine bird

In facultatively polygynous birds, secondary females of polygynously mated males typically have reduced annual reproductive success, because polygynous males provide less paternal care than monogamous males. Life history theory predicts that, as a result of increased reproductive investment, secondary females should suffer from reduced survival and lifetime reproductive success, but previous studies provided only weak support for this hypothesis. We used 7 years of data to study the fitness of female collared flycatchers Ficedula albicollis in relation to mating status by estimating survival and lifetime reproductive success. Taking differences in recapture probability into account, a mark-recapture analysis revealed that females observed at least once to breed as secondary female had higher survival than other females. This relationship was not confounded by laying date, because when we assessed the impact of laying date on survival, we found similar survival patterns. Females of polygynous males had reduced breeding success in terms of number of young fledged during the current reproductive event. However, during their lifetime females found at least once in primary or secondary mating status produced significantly more eggs, and at least the same number of fledglings and recruits as monogamous females. Thus, in the collared flycatcher, females of polygynously mated males seem to suffer from mating status during the most recent reproductive event, but considering survival and lifetime reproductive success, the apparently disadvantageous mating event is not necessarily associated with reduced residual reproductive value.     

Phylogeography and cryptic introduction of the ragworm Hediste diversicolor (Annelida,Nereididae) in the Northwest Atlantic

Many benthic marine invertebrates show striking range disjunctions across broad spatial scales. Without direct evidence for endemism or introduction, these species remain cryptogenic. The common ragworm Hediste diversicolor plays a pivotal role in sedimentary littoral ecosystems of the North Atlantic as an abundant prey item and ecosystem engineer, but exhibits a restricted dispersal capacity that may limit connectivity at both evolutionary and ecological time scales. In Europe, H. diversicolor is subdivided into cryptic taxa and genetic lineages whose distributions have been modified by recent invasions. Its origin in the northwest Atlantic has not been adequately addressed. To trace the age and origin of North American ragworm populations, we analyzed mtDNA sequence data (COI) from the Gulf of Maine and Bay of Fundy (n=73 individuals) and compared our findings with published data from the northeast Atlantic. Our results together with previous data indicate that two species of the H. diversicolor complex have independently colonized the northwest Atlantic at least three different times, resulting in two distinct conspecific assemblages in the Bay of Fundy and Gulf of Maine (respectively) that are different from the species found in the Gulf of St. Lawrence. North American populations had significantly lower genetic diversity compared with populations in the northeast Atlantic, and based on patterns of shared identity, populations in the Bay of Fundy originated from the Baltic Sea and North Sea. Populations from the Gulf of Maine were phylogenetically distinct and most likely originated from unsampled European populations. Analyses of the North American populations revealed patterns of post‐colonization gene flow among populations within the Gulf of Maine and Bay of Fundy. However, we failed to detect shared haplotypes between the two regions, and this pattern of complete isolation corroborates a strong phylogeographic break observed in other species.     

Reproductive success and pollen dispersal in urban populations of an insect-pollinated hay-meadow herb

Urban development increasingly occupies more landscape and is responsible for habitat alteration and fragmentation of so many ecosystems that urban environments can no longer be excluded from the areas to be considered in conservation strategies. Appropriate management measures ensuring population viability are needed, but due to the usually irreversible feature of the urbanisation process, the measures may differ from non-urbanised areas. Sufficient gene flow levels, leading to successful seed production, appear to be a key element in the sustainable persistence of insect-pollinated plant remnant populations. This study investigated reproductive success and pollen dispersal patterns in Centaurea jacea, an insect-pollinated hay-meadow herb, in the Brussels urban area (Belgium). Reproductive performance of urban park populations was compared with populations occurring in urban semi-natural sites. Pollen dispersal patterns were examined within and among-population patches from two urban parks using fluorescent dye as a pollen analogue. Urban park populations showed a higher reproductive success than those in urban semi-natural sites. Depositions of fluorescent dye particles on recipient individuals were found within populations (<20 m), among populations within parks (over a few hundred meters) and among parks (maximal investigated distance: 2.6 km), indicating pollinator movements and a potential gene flow by pollen over relatively long distances. Dye dispersal showed an exponential decay distribution with respect to spatial distance. The very small populations received more immigrant dye deposition when located at short distances from the source population. These findings demonstrate that urban populations do not perform worse in reproductive success than populations from agricultural landscapes, despite their small size, and can function as a network exchanging pollen. The conversion of urban park lawns into interconnected networks of ecologically managed hay meadows, allowing simultaneous flowering and seed production in many patches of target species, is to be strongly encouraged.     

Oxidative damage in relation to a female plumage badge: evidence for signalling costs

Badges of status may be controlled by costs derived from increased aggression from dominant individuals. This cost could be translated into elevated metabolic levels and a concomitant disruption of oxidative balance. Some females in Iberian pied flycatcher Ficedula hypoleuca populations exhibit a white forehead patch similar to that exhibited by all males in this species, functioning in aggressive interactions between females when competing for breeding sites. To test if social stress imposes costs on signalling, we painted white patches on females without natural patches (NP) and compared them with females with natural control (NU). We also over-painted the natural patch in other females (FP) and compared to females with control natural patches (FU). We obtained for the whole sample of females data on reproductive investment, morphology and oxidative damage measured by blood malondialdehydes (MDA), and in a subsample of females variables related to parental care during incubation and the early nestling stage. FP and FU did not differ significantly in any variable which negates an effect of paint itself. However, NP females showed significant higher levels of MDA than NU females when controlling for breeding success for the whole sample, and for female incubation attendance for the parental care subsample. When including the four treatments, there was a significant interaction between the paint treatment and the presence/absence of badges before the experiment when controlling for the significant negative effect of incubation attendance on MDA. Addition of a badge to females without one leads to increased oxidative damage possibly mediated by social control. Badges of status in female pied flycatchers may operate as badges of oxidative status.     

Gaping displays reveal and amplify a mechanically based index of weapon performance

Physical prowess, a key determinant of fight outcomes, is contingent on whole-organism performance traits. The advertisement of performance, via display, is poorly understood because it is unclear how information about performance is encoded into display characteristics. Previous studies have shown that weapon performance (i.e., bite force) predicts dominance and reproductive success in male lizards. We tested the hypothesis that gaping displays by adult male collared lizards (Crotaphytus) can provide an index of weapon performance by exposing the major jaw-adductor muscle complex and that white patches at the mouth corners amplify this index. For territorial adult males, the breadth of the muscle complex, which is not correlated with body size, was a strong predictor of bite force. For nonterritorial yearling males and females, however, measures of body and head size predicted bite force. The patches are highly conspicuous, exhibit UV-reflecting properties within the visual range of lizards, and provide size-independent information about bite force only in adult males. We conclude that exposure of the muscle complex during gaping displays can provide rival males with a reliable, body-size independent, biomechanically based index of weapon performance, an index that the mouth-corner patches amplify. Indexes that transmit information through mechanistic links to performance are expected to be widespread among animals.