Carry‐over effects of conditions at the wintering grounds on breeding plumage signals in a migratory bird: roles of phenotypic plasticity and selection

To understand the consequences of ever‐changing environment on the dynamics of phenotypic traits, distinguishing between selection processes and individual plasticity is crucial. We examined individual consistency/plasticity in several male secondary sexual traits expressed during the breeding season (white wing and forehead patch size, UV reflectance of white wing patch and dorsal melanin coloration) in a migratory pied flycatcher (Ficedula hypoleuca) population over an 11‐year period. Furthermore, we studied carry‐over effects of three environmental variables (NAO, a climatic index; NDVI, a vegetation index; and rainfall) at the wintering grounds (during prebreeding moult) on the expression of these breeding plumage traits of pied flycatcher males at individual and population levels. Whereas NAO correlates negatively with moisture in West Africa, NDVI correlates positively with primary production. Forehead patch size and melanin coloration were highly consistent within individuals among years, whereas the consistency of the other two traits was moderate. Wing patch size decreased with higher NAO and increased with higher rainfall and NDVI at the individual level. Interestingly, small‐patched males suffered lower survival during high NAO winters than large‐patched males, and vice versa during low NAO winters. These counteracting processes meant that the individual‐level change was masked at the population level where no relationship was found. Our results provide a good example of how variation in the phenotypic composition of a natural population can be a result of both environment‐dependent individual plasticity and short‐term microevolution. Moreover, when plasticity and viability selection operate simultaneously, their impacts on population composition may not be evident.     

Male territoriality and ‘sex confusion’ in recently adapted lizards at White Sands

The evolution of intersexual interactions, like mate choice, during ecological speciation has received widespread attention. However, changes in intrasexual interactions, like male territoriality, during ecological divergence are largely unexamined. We conducted field experiments with adaptively diverged populations of the eastern fence lizard (Sceloporus undulatus) to determine whether territorial males behaved differently towards ecologically similar vs. dissimilar intruders. We performed trials with light‐coloured males from White Sands, New Mexico and dark‐coloured males from the surrounding desert. We found that intruders from White Sands elicited more aggression than intruders from dark‐soil habitat. We also documented a case of ‘sex confusion’ where white‐sand males courted dark‐soil intruders. We found population differences in signalling patch size that can explain both aggression bias and sex misidentification. We argue that direct selection (for population recognition or optimal signal transmission) and indirect selection (by‐products of ecological adaptation) should influence both intersexual and intrasexual interactions during ecological speciation.     

Acrossocheilus spinifer, a new species of barred cyprinid fish from south China (Pisces: Teleostei)

Acrossocheilus spinifer sp. nov. is described from the river basins in Fujian Province and the Han Jiang basin in Guangdong Province, south China. It is one of the barred Acrossocheilus species sharing five or six vertical bars on the flanks, with each bar being two scales in width. Acrossocheilus spinifer sp. nov. is morphologically most similar to Acrossocheilus wenchowensis , but distinguished from it by: a colour pattern of all vertical bars extending ventrally to the second scale row below the lateral line, and having the second bar placed posterior to the base of the last simple dorsal‐fin ray, a blunt snout; and a stout, last simple dorsal‐fin ray. Acrossocheilus spinifer sp. nov. resembles Acrossocheilus kreyenbergii and Acrossocheilus stenotaeniatus with which it shares a stout, last simple dorsal‐fin ray with a serrated posterior edge, but is separated from both species by having a narrow median interruption in the lower lip, a lower jaw fully covered by the lower lip when viewed ventrally and the second vertical bar placed posterior to the base of the last simple dorsal‐fin ray. The absence of a longitudinal stripe extending along the lateral line on the flanks in males readily distinguishes A. spinifer sp. nov. from four other species, namely Acrossocheilus fasciatus , Acrossocheilus paradoxus, Acrossocheilus parallens , and Acrossocheilus jishouensis .     

It is all in the head: morphological basis for differences in bite force among colour morphs of the Dalmatian wall lizard

Males of the lizard Podarcis melisellensis occur in three distinct colours that differ in bite performance, with orange males biting harder than white or yellow ones. Differences in bite force among colour morphs are best explained by differences in head height, suggesting underlying variation in cranial shape and/or the size of the jaw adductors. To explore this issue further, we examined variation in cranial shape, using geometric morphometric techniques. Additionally, we quantified differences in jaw adductor muscle mass. No significant differences in size corrected head shape were found, although some shape trends could be detected between the colour morphs. Orange males have relatively larger jaw adductors than yellow males. Not only the mass of the external jaw adductors, but also that of the internal jaw adductors was greater for the orange morph. Data for other cranial muscles not related to biting suggest that this is not the consequence of an overall increase in robustness in orange individuals. These results suggest that differences in bite performance among morphs are caused specifically by an increase in the mass of the jaw adductor, which may be induced by differences in circulating hormone levels.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 13–22.     

Sex‐related spatial patterns of Poa ligularis in relation to shrub patch occurrence in northern Patagonia

Abstract. Poa ligularis is a dioecious species and a valuable forage plant which is widespread in the arid steppe of northern Patagonia (Argentina). The vegetation in these areas consists of a system of perennial plant patches alternating with bare soil areas defining contrasting micro‐environments. We hypothesized that (1) male and female individuals of P. ligularis are spatially segregated in different micro‐environments, (2) the intensity of spatial segregation of sexes depends on plant structure and (3) spatial segregation of sexes is enhanced by competitive interactions between the sexes within the vegetation patches. We analysed the spatial distribution of female and male individuals in relation to the spatial pattern of vegetation in two areas differing in their vegetation structure. The location of P. ligularis within patches where either male, female or both sexes occurred was also analysed. The results indicate that different patterns of spatial distribution of sexes of P. ligularis may be found at the community level depending on the dominant life forms and geometric structure of plant patches. Where patches are of a lower height, with a high internal patch cover, individuals of both sexes are concentrated within patch canopies. In sites characterized by large, tall patches and less internal patch cover suitable microsites for female and male P. ligularis occur both within and outside the patch with males located at further distances from the patch edge. Where the patch is large and tall enough to allow the establishment of males and females at relatively high numbers, males occupy the patch periphery or even colonize the interpatch bare soil. These spatial patterns are consistent with selective traits in which females better tolerate intraspecific competition than males, while males tolerate wider fluctuations in the physical environment (soil moisture, nitrogen availability, wind intensity, etc.).     

Forehead Patch Size Predicts the Outcome of Male–Male Competition in the Pied Flycatcher

Males of many animal species express ornaments that affect their reproduction opportunities through male–male competition or female mate choice. Such ornaments can, for example, inform conspecifics about the fighting ability, condition or territory ownership of the bearer. Pied flycatcher (Ficedula hypoleuca) males have a conspicuous white forehead patch that varies greatly in size. We examined whether the white forehead patch is an intrasexually selected trait in a Finnish population. We artificially manipulated forehead patch size to represent two naturally occurring extremes and competed males against each other in the presence of a female. Males with a large forehead patch were more aggressive than males with a small patch, whereas the original patch size of the male had no influence on aggression. Neither manipulated nor original patch size influenced resource dominance (over female or nest box). These results indicate that forehead patch signals fighting ability of the bearer in the pied flycatcher. The next step is to find out what kind of costs may maintain the honesty of this signal.     

Evidence for sexual dichromatisms in spawning aggregations of yellowfin grouper Mycteroperca venenosa and tiger grouper Mycteroperca tigris from the southern Gulf of Mexico

Colour pattern characteristics and gonad histology were used to detect sexual dichromatisms in yellowfin grouper Mycteroperca venenosa and tiger grouper Mycteroperca tigris from the Campeche Bank, Mexico. Specimens were obtained from commercial catches between March and May during 2002 and 2004. All specimens were examined dead. Ninety-seven per cent of males had different sex-associated colour patterns. Male yellowfin grouper displayed a bright yellow blotch on both sides of the lower jaw while females retained a reddish lower jaw. Male tiger grouper had uniform dark pectoral fins while females had bright orange pectoral fins. In situ observations of live fishes at fishing sites showed the lower jaw and pectoral fin colourations to be clearly visible underwater at a depth of 35 m. All males of both species and most females (80% yellowfin grouper and 98% tiger grouper) were sexually active and probably caught during their spawning season. This suggests that distinct colourations observed for male M. venenosa and M. tigris may be seasonal displays associated with spawning. Both the lower jaw and pectoral fin colourations were still visible in dead fishes after several days on ice. Differences observed for ray length of exserted vertical fins in tiger grouper specimens were probably not a sex-associated characteristic.     

A New Species of Leptolalax （Anura： Megophryidae） from Southern China

A new species, Leptolalax laui sp. nov. is described based on specimens collected from Hong Kong and Shenzhen City, Guangdong Province, China. The new species can be distinguished from other known congeners by morphological and molecular data. The new species is characterized by the following characters： 1） small size （adult males SVL 24.8.1 mm-26.7 mm）; 2） near immaculate creamy white chest and belly; 3） broad lateral fringes on toes; 4） head longer or as long as wide; 5） distinct dark brown spots in flank; 6） moderate dermal fringes on fingers; 7） brown or reddish-brown dorsum with fine round scattered tubercles; 8） thin traverse brownish-grey bars on the dorsal surface of tibia and lower arms; 9） longitudinal ridges under toes not interrupted at the articulations.     

POPULATION DIVERGENCE IN SEXUAL ORNAMENTS: THE WHITE FOREHEAD PATCH OF NORWEGIAN PIED FLYCATCHERS IS SMALL AND UNSEXY

Models of sexual selection suggest that populations may easily diverge in male secondary sexual characters. Studies of a Spanish population of the pied flycatcher, Ficedula hypoleuca, and a Swedish population of the closely related collared flycatcher, F. albicollis, have indicated that the white forehead patch of males is a sexually selected trait. We studied the white forehead patch of male pied flycatchers (n = 487) in a Norwegian population over seven years. Males with large forehead patches were in general more brightly colored, but patch height was not correlated to body mass, body size, or parasite loads. Conditions during the nestling period did not seem to influence patch height as an adult. Patch height increased slightly from the first to the second year as adults, but then remained relatively constant at higher ages. Patch height was not related to survival. Year-to-year changes showed that males who increased in patch height also increased in body mass, suggesting that expression of the forehead patch may be partly condition dependent. However, changes in body mass explained only a small proportion of the variance in patch height between males. Thus, patch height would not be a good indicator of male quality. Furthermore, patch size was also not related to male ability to feed nestlings, indicating that females would not obtain direct benefits by choosing males with large patches. However, patch height could be a Fisher trait, but this requires heritability and there was no significant father-son resemblance in patch height. Comparisons of the males visited by each female during the mate sampling period indicated that chosen males did not have larger forehead patches than rejected males. Experimental manipulation of patch height did not affect male mating success. These results indicate that females do not use patch size as a mate choice cue. Finally, patch height did not predict the outcome of male contests for nestboxes, suggesting that the forehead patch is not an intrasexually selected cue of status. Norwegian pied flycatchers have smaller forehead patches than both Spanish pied flycatchers and Swedish collared flycatchers. We suggest that this pattern may be explained by the lack of sexual selection on the forehead patch in the Norwegian population as compared to the other populations, where the patch is apparently sexually selected. We discuss possible reasons for these population divergences, such as female choice on an alternative secondary sexual character (general plumage color) and speciation among Ficedula flycatchers.     

Morphological evidence for the biological role of caniniform teeth in combtooth blennies (Blenniidae, Teleostei)

Combtooth blennies have recurved, fang-like caniniform teeth at the rear end of a single row of incisiform teeth. The lengths and positions of these canines were measured in the lower jaws of males of 14 species of Mediterranean Blenniidae. In four species, lower jaw canines were measured in males and females, while in one species, the upper jaw canines of both sexes were also measured. Relative (to body length) canine length in males tends to be significantly greater (10–40%) than in females. There are significant interspecific differences in relative canine length, with smaller species tending to have relatively larger teeth. No significant correlation was obtained between canine length and importance of animal prey in the diet, nor with 'hole fit' of males, which may be related with the intensity of paternal care. We suggest that canines in combtooth blennies are predominately used for predator deterrence and agonistic interactions.     

Free imidazole compounds in white and dark muscles of migratory marine fish

1. Extractive nitrogenous components were analyzed in specimens of white and dark muscles of three tunas and the Pacific saury. 2. Tunas contained high concentrations of histidine, anserine and creatine in white muscle, and of taurine, anserine and creatine in dark muscle. 3. Imidazole compounds were determined in both muscles of 11 migratory marine fish. Histidine was found in great quantities in all species except swordfish, anserine was found in relatively large amounts in tunas and swordfish, but carnosine was only present in small amounts in yellowfin and skipjack tunas. 4. The dark muscle had a lower content of imidazole compounds than white muscle.     

A new species of the clingfish genus <Emphasis Type="Italic">Apletodon</Emphasis> (Teleostei: Gobiesocidae) from Sao Tome and Principe,Eastern Central Atlantic

The clingfish Apletodon wirtzi sp. nov. is described on the basis of ten specimens and colour photos from Bombom Island, Principe Group, Sao Tome and Principe, eastern central Atlantic Ocean. The species is very small, apparently not exceeding 16 mm total length; it is characterized by having three pores in the mandibular canal, the head length 2.2–2.5 in standard length: SL, the head broad, head width in males 3.6–4.0 (mean 3.8) in SL, the snout in males long, more or less pointed, conical, preorbital length in males 3.1–4.0 in head length, the occipital region with a large pinkish blotch behind each eye (in alcohol specimens), and the lower sides of the body with a row of dark blotches, scattered with white spots in between. The new species is compared with other species of the genus; a key to the males of the four known species of the eastern Atlantic genus Apletodon is presented. Supplementary material to this paper is available in electronic format at     

Different roles of feeding and protection in diel microhabitat choice of sexes in<Emphasis Type="Italic"> Idotea baltica</Emphasis>

In Idotea baltica, a marine isopod that lives and feeds on the brown alga Fucus vesiculosus, microhabitat choice differs between sexes so that males are found more often than females on the light-coloured and exposed apical parts of the alga. We investigated how the requirements of avoiding visual predators and feeding were related to microhabitat choice in relation to diurnal and life-cycle stage in males and females. Faced with a choice between an apical and a basal piece of the alga, females spent more time than males on the basal piece, but this difference was not due to food choice. Faced with a choice between a dark, concealing and a light, exposing background, the preference for a dark background was stronger at day than at night, and stronger in females than in males. This suggests that a sex difference in the importance of avoiding visual predators can explain the sex difference in microhabitat choice. Further, the preference for a dark background and night feeding both increased with age, suggesting that feeding is increasingly subordinated to the need to avoid visual predators. Our experiment found no effect of the presence of the opposite sex on microhabitat choice. Our results support the hypothesis that the sexes trade off feeding against predation risk differently, presumably because growth is more important to males than to females, which have more to gain by protection and therefore spend more time on the lower parts of the alga. Received: 22 March 1999 / Accepted: 7 October 1999     

Revision of the wobbegong genus <Emphasis Type="Italic">Orectolobus</Emphasis> from Japan,with a redescription of <Emphasis Type="Italic">Orectolobus japonicus</Emphasis> (Elasmobranchii: Orectolobiformes)

The wobbegong genus Orectolobus occurring from Japan and its adjacent waters was reviewed, and O. japonicus was redescribed using ten specimens including a syntype. Only one species, O. japonicus, was regarded as valid among three species previously recorded from Japan. Orectolobus maculatus had been erroneously described from Japan because of the past nomenclatural confusion with O. japonicus. The specimen called “Karakusa-oose,” which had been identified with O. ornatus, was regarded as an irregular form of O. japonicus. Orectolobus japonicus was distinguished from the other congeners by having no tubercles on the body, five to eight dermal lobes in the preorbital group, two distally notched lobes in the postspiracular group, no dermal lobes on lower jaw, nasal barbel with a branch, no supraocular knob, relatively high dorsal fins with usually concave posterior margins, a lower number of precaudal vertebrae and intestinal valve turns, saddles on the back without black borders and yellowish and variously shaped blotches, not forming white O-shaped spots.     

Geographic variation in the carotenoid plumage pigmentation of male house finches (Carpodacus mexicanus)

Geographic variation in both the colour and pattern of carotenoid plumage pigmentation displayed by males in two subspecies of house finches ( Carpodacus mexicanus frontalis and C. m. griscom ) was quantified. The extent of ventral carotenoid pigmentation (patch size) differed markedly between these two subspecies; frontalis males from the U.S. (New York, Michigan, California and Hawaii) displayed a medium patch extending from their throats to their lower bellies, while griscomi males sampled in Guerrero, Mexico displayed small patches restricted to their throats. Frontalis males sampled in Michigan and New York and griscomi males were relatively bright in colouration, while frontalis males sampled in Hawaii were relatively drab. Populations of frontalis in California showed substantial local variation in average male colouration: in two areas only 12 km apart males were as colourful and as drab as any population sampled. In aviary experiments in which they were fed either a plain seed diet or a diet supplemented with red carotenoid pigments during moult, males from all populations converged on a similar appearance, except that griscomi males attained a brighter plumage than frontalis males when their diet was supplemented with red pigments. Regardless of diet, the difference in patch size between frontalis and griscomi males persisted after moult in captivity. The author concludes that the difference in patch size between frontalis and griscomi males reflects genetic differences between these populations, but that the differences in the mean plumage colouration of males among populations reflect differences in the access that males have to carotenoid pigments during moult.     

Variation of mandible size in red deer<Emphasis Type="Italic">Cervus elaphus hispanicus</Emphasis> from southern Spain

An osteometric study was carried out on 126 mandibles of male and female deerCervus elaphus hispanicus Linnaeus, 1758 hunted between 1992 and 1998 in Sierra Morena (SE Spain). Age of animals was estimated in months using thin sections of the first lower molar (M1). Length measurements of the mandible were plotted against age. An analysis of variance was used to assess the influence of year of birth, sex, size of estates and deer abundance within each closed estate on mandible length. Results reveal a differential growth between males and females, along with differences in jaw morphology. Growth is complete at 55–56 months (4.5–5 years) in females and at 80–85 months (6.5–7 years) in males. The jaw angle, the condylar process and the coronoid process are larger in males. All measurements were larger in animals born in 1989 or earlier, and lower in animals born after 1993, revealing the strong influence of climatic conditions. A negative effect of the deer abundance on growth was also observed.     

Distribution and reproductive success of the Pied Flycatcher Ficedula hypoleuca in relation to forest patch size and vegetation characteristics; the effect of scale

We studied habitat choice, distribution and reproductive success in the Pied Flycatcher Ficedula hypoleuca in relation to the spatial structure of a fragmented forest area. Two hierarchical levels of forest habitat structure were used: (1) forest patch size (macrohabitat) and (2) vegetative structure within forest patches (microhabitat). In spring, both males and females settled preferentially in large and medium-sized forest stands (>1 ha) where breeding density was also higher than in small stands (<1 ha). Stands <5 ha were occupied later, and the proportion of unpaired males was higher there than in large stands (>5 ha). The known age distributions of breeding birds and breeding success were independent of forest patch size. Nest predation rate was not associated with stand size or nest distance from the forest edge. The preference of breeding birds for large forest patches was presumably related to the higher amount of resources (nest sites, mates, food) a large patch can offer for reproduction compared with a small patch. At the microhabitat level, territories of old males were characterized by relatively more deciduous trees that contained more invertebrate food than coniferous tree-dominated territories of yearling males. The reproductive success of old males, as measured by the number of fledged young per male, was higher than that of yearling males. This suggests that the larger body size and blacker plumage of old males possibly contributed to male dominance and that old males excluded younger males from preferred habitats through territorial behaviour. Our results suggest that habitat choice of the Pied Flycatcher was affected by both micro- and macrohabitat. On the basis of settlement pattern, density and reproductive success, the distribution of Pied Flycatchers across forest stands of different sizes followed the ideal-free distribution model, whereas at the microhabitat level, age-related unequal distribution of males followed the ideal-despotic model of Fretwell and Lucas.     

Females Paired with More Attractive Males Show Reduced Oxidative Damage: Possible Direct Benefits of Mate Choice in Pied Flycatchers

Direct benefits of female mate choice may concern female fertility and fecundity but also physiological status. In birds with biparental care, males may contribute to improve the condition and health of their pair‐mates through help in constructing nests, incubation or incubation feeding and nestling provisioning. They may also reduce harassment of females by non‐pair males. A consequence of these male activities could be expressed in terms of oxidative damage, which may depend on metabolic effort and social stress. Here, we have related male contribution to parental and territorial duties to female oxidative status in the pied flycatcher Ficedula hypoleuca, a species where preferred males present darker dorsal plumage and, in Iberian populations, a large white forehead patch. Darker males were paired with females with high incubation attendance and reduced nestling provisioning rates, which may lead to reduced female exertion. These males owned nest boxes at which there were fewer visits by non‐pair males. Although females paired with dark mates worked less hard, they were able to raise more fledglings. Female oxidative damage measured as malondialdehyde (MDA) level in plasma declined with increasing incubation attendance and male incubation feeding. Moreover, levels of MDA in females declined with both darkness of male dorsal plumage and male forehead patch size when controlling for female forehead patch size and male age. The effect of male plumage darkness was especially strong. Females paired with middle‐aged males (2–3 yr) showed reduced levels of MDA compared with those paired with 1‐yr‐old and more than 3‐yr‐old males. Male age could not explain the effects of male attractiveness. Females paired with attractive males were more successful in reproduction while suffering reduced oxidative damage, possibly mediated by help during incubation and nestling rearing from their pair‐mates. Although correlative, the evidence suggests direct benefits of females paired with more attractive males.