Plumage Research Articles

New feather mite taxa of the Rhytidelasma Gaud, 1966 generic group (Astigmata: Pterolichidae) from the red-flanked lorikeet Charmosyna placentis placentis (Psittacidae).

Two new genera and species related to Rhytidelasma Gaud, 1966 and Lorilichus Atyeo & Gaud, 1991 are described from the plumage of the red-flanked Lorikeet Charmosyna placentis placentis (Temminck) (Psittacidae). These are Ceratolichus mirabilis n. g., n. sp. and Charmosylichus anamphiestos n. g., n. sp. In addition, an improved diagnosis of Lorilichus is proposed and two new species of this genus are described from the red-flanked lorikeet, L. longiphallos n. sp. and L. paralobiger n. sp.

Genetic divergence in the superspecies Manacus

The bearded manakins in the genus Manacus are lekking, neotropical passerines. Male plumage colour varies with geographical location and classification is based solely on these plumage patterns. It has recently been suggested that in this group of birds, plumage patterns may be a misleading taxonomic character. In this study we used microsatellite variation in a collection of museum samples to establish the amount of genetic divergence between the previously described bearded manakin species/subspecies. We found substantial genetic substructuring between species/subspecies and that plumage patterns indeed may be a misleading taxonomic character because the presence of yellow in male nuptial plumage is found in most divergent forms. We did not detect a significant isolation by distance relationship although the P-value was close to significance. Physical barriers such as rivers and mountains may affect gene flow and play a role in shaping genetic structure of the genus Manacus. Accordingly, boundaries between species/subspecies often coincide with large rivers, mountains and seas.

Molt and Plumage Terminology of Howell et al. (2003) Still May not Reflect Homologies

Abstract Forty-five years ago Humphrey and Parkes (1959) introduced their system of nomenclature for molts and plumages (the H-P system). They claimed that their biologically neutral terminology, independent of the bird's life cycle events, could accurately reflect homologies of molts and plumages across a wide range of avian taxa. Soon, however, several investigators reported trouble adapting the terminology to unusual sequences of molts and plumages, and others expressed doubts that it could accurately reflect true phylogenetic homologies. Howell et al. (2003) reviewed these problems and proposed a modification of the original H-P system that identifies the juvenal plumage as the first basic plumage, and any postjuvenal plumages that are not repeated in later plumage cycles as formative plumages. By doing this, they solved the problem of adapting the original H-P system to plumages of the first year of life in many species. However, they have not overcome the arguments of Stresemann (1963) and Amadon (1966) that patterns of molting and plumage are so variable, and so subject to natural selection, that there is doubt that the H-P terminology can reveal phylogenetic homologies even among closely related species. Molting and plumage sequences within a clade comprising four species of the genus Carduelis confirm that within this limited grouping, molting and plumages are highly variable, and apparent homologies are not revealed in the H-P terminology as modified by Howell et al. La Terminología de la Muda y el Plumaje de Howell et al. (2003) Podría no Reflejar Aún las Homologías Resumen. Hace cuarenta y cinco años, Humphrey y Parkes (1959) introdujeron su sistema de nomenclatura para la muda y el plumaje (el sistema H-P). Ellos sugirieron que su terminología biológicamente neutra, independiente de los eventos del ciclo de vida de las aves, podría reflejar con exactitud las homologías de la muda y el plumaje a través de un amplio espectro de taxa. Sin embargo, poco después varios investigadores notaron problemas al adaptar la terminología a secuencias inusuales de muda y plumajes, y otros expresaron dudas en cuanto a si podría reflejar con exactitud las verdaderas homologías filogenéticas. Howell et al. (2003) revisaron estos problemas y propusieron una modificación del sistema H-P original que identifica el plumaje juvenil como el primer plumaje básico, y cualquier plumaje post-juvenil que no se repita en ciclos posteriores del plumaje como plumaje formativo. Haciendo esto, solucionaron el problema de adaptar el sistema H-P original a plumajes del primer año de vida de muchas especies. Sin embargo, no lograron resolver las críticas de Stresemann (1963) y Amadon (1966) de que los patrones de muda y plumaje son tan variables y tan propensos a ser afectados por selección natural, que es dudoso que la terminología H-P pueda revelar homologías filogenéticas, aún entre especies estrechamente relacionadas. Las secuencias de muda y plumaje al interior de un clado formado por cuatro especies del género Carduelis confirman que dentro de este limitado grupo, la muda y el plumaje son áltamente variables y que algunas homologías aparentes no son reveladas por la terminología H-P con las modificaciones de Howell et al.

Determining Evolutionary Homologies of Molts and Plumages: A Commentary on Howell et al. (2003)

AbstractHowell et al. (2003) argue that the Humphrey-Parkes (H-P) system of molt terminology is flawed because it requires using traditional first prebasic molt as the starting point for plumage succession that results in noncorrespondence between nomenclature and presumed homology in first basic plumages. However, the H-P system does not require this. Second, they argue that plumage color can be a misleading criterion for evaluating plumage homologies. I show, however, that the timing and extent of molts, and thus their homologies, can de documented more accurately by using plumage color than by not doing so. Howell et al. (2003) propose a revised H-P system. To follow their system, one must accept their notion that no first-cycle molts are homologous with prebasic molts in subsequent molt cycles. However, this is not so as many species have a molt in their first cycle that is homologous to definitive prebasic molt. In addition, Howell et al.'s (2003) system does not offer any new or better criteria for identifying homologies than those suggested by Humphrey and Parkes (1959) and, thus, is not an improvement on the H-P system. First-cycle molts and plumages of most birds are poorly known. Therefore, we will not have sufficient data to determine whether new molts have been evolutionarily added to the first cycle, as suggested by Howell et al. (2003), until the molts of many more species of birds are studied. Further, these studies must be done on closely related species, not phylogenetically distant ones as proposed by Howell et al. (2003).Determinación de las Homologías Evolutivas de la Muda y el Plumaje: Un Comentario sobre Howell et al. (2003)Resumen. Howell et al. (2003) aducen que el sistema Humphrey-Parkes (H-P) de terminología para la muda es erróneo porque requiere utilizar la primera muda prebásica como el punto de partida para la sucesión del plumaje, lo que resulta en falta de correspondencia entre la nomenclatura y las presuntas homologías en los primeros plumajes básicos. Sin embargo, el sistema H-P no requiere esto. Segundo, ellos argumentan que el color de plumaje puede ser un criterio engañoso para evaluar las homologías del plumaje, pero yo demuestro que el momento y la extensión de las mudas, y por tanto sus homologías, pueden documentarse con mayor exactitud utilizando el color del plumaje que no haciéndolo. Howell et al. (2003) proponen un sistema H-P revisado que implica aceptar su noción de que ninguna de las mudas del primer ciclo es homóloga con mudas prebásicas de ciclos de muda subsiguientes. Sin embargo, esto no es así, pues muchas especies tienen una muda en su primer ciclo que es homóloga a la muda prebásica definitiva. Adicionalmente, el sistema de Howell et al. (2003) no ofrece criterios nuevos o mejores para identificar las homologías que aquellos sugeridos por Humphrey and Parkes (1959), por lo que no representa un mejoramiento del sistema H-P. Las mudas y los plumajes del primer ciclo de la mayoría de las aves son poco conocidos. Por lo tanto, hasta que no se estudie la muda en muchas más especies de aves, no tendremos suficientes datos para determinar si nuevas mudas se han adicionado evolutivamente al primer ciclo como Howell et al. (2003) sugirieron. Más aún, dichos estudios deben hacerse en especies estrechamente relacionadas, no en aquellas filogenéticamente distantes como Howell et al. (2003) propusieron.

Do body condition and plumage during fuelling predict northwards departure dates of Great Knots Calidris tenuirostris from north‐west Australia?

It is often assumed that strong selection pressures give rise to trade‐offs between body condition and time in long‐distance migrating birds. Birds that are ‘behind schedule’ in fuel deposition or moult should delay departure, and this should result in a negative correlation between initial condition and departure date. We tested this hypothesis in the Great Knot Calidris tenuirostris migrating from north‐west Australia to eastern Asia en route to Siberia. Great Knot gain mass and moult into breeding plumage before leaving northern Australia in late March and early April, and fly 5400–6000 km to eastern China and Korea. We radiotracked 27 individuals (17 males and ten females) to determine departure dates; 23 migrated and four remained in Australia. We characterized body condition at capture using body mass, predicted pectoral muscle mass (based on ultrasound estimates of the size of the pectoral muscles) and breeding plumage scores. Residual condition indices were uncorrelated, indicating that at the individual level, variation in one fuelling component was not strongly associated with variation in the other components. Birds that did not depart had lower residual body mass and breeding plumage indices than those that did migrate; these four birds may have been subadults. Neither sex, size nor the condition indices explained variation in departure date of migrants. Reasons for this are explored. Departure dates for northward migrating waders indicate that the migration window (span over which birds depart) decreases with proximity to the northern breeding grounds. We suggest that migration schedules become tighter as birds get nearer to the breeding grounds. Thus the lack of a relationship between condition and departure date in Great Knots may reflect the fact that the departure episode under study is the first one in sequence and is still 4–8 weeks before breeding.

Ground jays expand plumage to make themselves less conspicuous

Ground jays expand plumage to make themselves less conspicuous

Pay‐offs and penalties of competing migratory schedules

We relate variation in the timing of arrival by migrating birds breeding at northerly latitudes to individual differences in the prior accumulation of energy stores. Balancing starvation risks early in the season against the almost universal declining trend in reproductive prospects with advancing date is seen as an individual decision with fitness consequences. We review three studies implicating events at the staging sites or in winter in setting the individual migratory schedule. Climate change influences the timetable of a pied flycatcher (Ficedula hypoleuca) population breeding in The Netherlands and wintering in West Africa, followed since 1960. Mean air temperature in the period mid April‐mid May (arrival and laying) increased and laying date advanced by 10 days. Still, in recent years most birds did not lay early enough to maximise fitness (determined by recruitment and parental survival) whereas many parents achieved this goal in 1980–1985. As the flycatchers have not started to arrive earlier, some ecological constraint further upstream is postulated (possibly the hurdle of the crossing of Sahara and Mediterranean). The ability to follow individual migrants provides a second avenue to assess the fitness implications of migratory schedules. Thus, brightly coloured male bar‐tailed godwits (Limosa lapponica) captured in the Dutch Wadden Sea (the intermediate staging site linking a West African wintering area with breeding sites in arctic Russia) and traced with miniature radio‐transmitters did not depart early. The ‘best’ males (with bright breeding plumage) were picked up by the listening stations in Sweden 650 km further along the migratory route ten days later than the paler individuals. If early arrival confers the competitive advantage of prior occupancy but increases mortality, the ‘best’ males may be able to afford arriving later and thus avoid some of the survival costs. Return rate of the ‘bright’ males to the staging site in later seasons was indeed higher than for the ‘pale’, early males. Intensive observation of pink‐footed geese (Anser brachyrhynchus) fitted with coded neck‐collars substantiate the tight relationship between energy stores (fat) accumulated up to final departure from the final staging site (Vesterålen, N. Norway) en route to the nesting grounds (Spitsbergen) and subsequent success. The breeding outcome of individual parents (accompanied by juveniles or not) could be related to observations of body condition before departure (visual ‘abdominal profile index’). Recently, perceived conflicts with agriculture have resulted in widespread harassment by humans. The geese have: drastically shortened their stay on the Vesterålen, fail to achieve the body condition usual a decade ago and reproductive output has fallen. Although the geese are currently pioneering new staging sites, an adequate alternative has not materialised, underlining the critical role of the final take‐off site.

The first record of Dunlin (Calidris alpina) in Paraguay and a summary of South American records of the species

A Dunlin (Calidris alpina) in full breeding plumage was discovered at Laguna Ganso in the Paraguayan Chaco on 29 June 2001. This bird represents the first record of the species for the country and for the interior of South America. Although primarily a species of higher northern latitudes, there are a number of previous records from coastal South America.

A postnuptial staging site for the Black Tern (Chlidonias niger) and White-winged Tern (C. leucopterus) in the Sivash, Ukraine

Numbers, biometrics and feeding ecology of two marsh tern species, the Black Tern (Chlidonias niger) and White-winged Tern (C.leucopterus) were studied in the Sivash, Ukraine in July–August 1996 and 1998. In 1998 the Sivash region was counted completely and, including the nearby Molochnyi Liman, an estimated 30,000 to 40,000 Black Terns and 50,000 to 60,000 White-winged Terns were present. This makes the Sivash region one of the most important postnuptial stopover sites for these marsh terns in Europe with respectively 15 % and 25 % of the total Eurasian population present at any one moment. Both species showed clear habitat segregation. Black Terns fed in deep brackish waters and White-winged Terns fed in terrestrial habitats as well as shallow fresh and brackish waters. Black Terns were feeding mainly on relatively large fish (of 6–12 cm), mostly on Big-scale Sand SmeltAtherina boyeri bonapartii. White-winged Terns were feeding on more diverse food types like large insects and small fish. In the Lucerne fields they were feeding on caterpillars and in the steppe areas on grasshoppers or spiders. In the very shallow lagoons they fed on young mullet but mainly in the evenings. Both species had high food intake rates as compared to terns feeding in other food-rich coastal wetlands. Both tern species moulted primaries as well as body feathers and they changed breeding plumage into non-breeding within three to four weeks. Even some juveniles changed their first body feathers into first winter plumage, an event unknown to date for both species in this period. The small sample of captured adult Black Terns showed a mass increase for active moulting terns resulting in higher body mass in later moult stages. The high body mass of marsh terns in August compared to the breeding season indicates fat storage prior to migration. The results support the theory that marsh terns are long non-stop migrants that use the Sivash as a last stopover before their flight to Africa.

Dietary and Sexual Correlates of Carotenoid Pigment Expression in Dove Plumage

AbstractCarotenoid pigmentation in birds' plumage is considered an honest indicator of phenotypic quality, and thus a target of sexual selection. But carotenoids also fulfill essential physiological functions, and therefore, carotenoids should only appear in plumage if they are in excess of those needed physiologically. We explored the presence of carotenoid-based plumage coloration in columbids and its association with diet and sexual dichromatism using a comparative analysis. We found that carotenoid plumage pigmentation appeared three times independently in doves, and that these events were always associated with frugivorous feeding habits. This suggests that expression of carotenoid-based plumage color in granivorous species may be constrained by the scarcity of carotenoids in their diet. However, more than half of the frugivorous species lack carotenoid-pigmented plumage, indicating that rich dietary sources of these compounds are a necessary but not sufficient cause for their expression in plumage. Analyzing 12 pairs of sister taxa, we found that plumage dichromatism was neither associated with the amount of carotenoid pigment present in the plumage nor with the sexual dimorphism in carotenoid-pigmented plumage. Although the presence of carotenoid-based plumage coloration has been related to sexual selection in several taxa, we failed to show such an association in columbids.Correlación de la Expresión de Pigmentos Carotenoides en el Plumaje de Palomas con la Dieta y la Selección SexualResumen. Los pigmentos carotenoides en el plumaje de las aves son considerados indicadores honestos de la calidad fenotípica y, por lo tanto, objetos de selección sexual. Sin embargo, los carotenoides también cumplen funciones fisiológicas esenciales, por lo cual aquellos que se expresan en el plumaje deberían estar en exceso de los utilizados a nivel fisiológico. Exploramos la presencia de carotenoides en el plumaje de las palomas y su asociación con la dieta y el dicromatismo sexual usando un análisis comparativo. Encontramos que el plumaje carotenoide apareció tres veces independientemente en palomas, y que estos eventos estaban siempre asociados a frugivoría. Esto sugiere que la expresión de carotenoides en el plumaje de especies granívoras puede estar restringida por la escasez de estos pigmentos en la dieta. Sin embargo, más de la mitad de las especies frugívoras carecen de plumaje carotenoide, indicando que alimentos ricos en este compuesto son una causa necesaria pero no suficiente para su expresión en el plumaje. Analizando 12 pares de taxa hermanos, encontramos que el dicromatismo no estaba asociado ni a la cantidad de pigmentos carotenoides presentes en el plumaje ni al dimorfismo sexual de plumaje carotenoide. Aunque la presencia de plumaje carotenoide ha sido relacionada con la selección sexual en numerosos grupos, no encontramos una asociación similar en palomas.

Female mate preferences and subsequent resistance to copulation in the mallard

In the majority of socially monogamous bird species, females solicit or accept copulations from males other than their partner. Females may gain direct benefits from extrapair males, such as greater access to resources, or indirect genetic benefits that will influence the future success of their offspring. However, one group of birds appears to be the exception to this general rule; in the wildfowl (Anseriformes), all extrapair copulations appear to be resisted by females. It has been suggested that resistance behavior may be a strategy to allow females a greater choice of mates, either at the precopulatory level (to promote choice of copulation partner) and/or the postcopulatory level (to promote multiple mating to increase their choice of sperm). This paper examines the function of female resistance behavior in one of the dabbling ducks, the mallard (Anas platyrhynchos). Observations on a marked population of wild mallard and experiments with captive birds found that although females showed a strong preference for particular males that are the first to molt into their breeding plumage, male attractiveness did not influence female responses to pair or extrapair copulation attempts. Female resistance decreased the likelihood that copulation attempts would end in successful insemination. The findings did not support the hypothesis that females resist copulations to promote female choice and the reasons why waterfowl may benefit from avoiding all extrapair copulations are discussed. Copyright 2003.

Differential moult patterns in relation to antipredator behaviour during incubation in four tundra plovers

Golden plovers and Grey Plovers Pluvialis spp. all have very distinct breeding plumage rich in contrast, with a conspicuous black belly and breast bordered by a bright white fringe. Eurasian Golden Plovers are known partly to replace their breeding plumage with striped yellow feathers during incubation, different from both breeding and non‐breeding plumages. In this study a similar partial breeding moult was observed in Pacific Golden Plovers and American Golden Plovers caught on the nest or collected during incubation, although the feathers did not differ clearly from those of non‐breeders. This moult starts during incubation and precedes the post‐breeding moult into non‐breeding plumage. Because the lighter feathers reduce the contrast between the black belly and the white flanks, we suggest that during incubation the plumage characteristic that plays an important role in mate choice is no longer important; at this stage it is better for the bird to be inconspicuous. Additional information on museum skins of golden plovers and of Grey Plovers indicated that only the three golden plovers undergo this partial moult, but that Grey Plovers in general retain full breeding plumage throughout incubation. The three golden plovers also resemble each other in their generally very passive nest defence strategies. In contrast, the larger Grey Plovers actively chase and attack aerial and ground predators. Thus, a reduced conspicuousness of the body plumage during incubation is likely to benefit the golden plovers more than the Grey Plover. We suggest that nest defence behaviour, plumage characteristics and perhaps size have co‐evolved as a response to different selection pressures in golden plovers and Grey Plover, but alternative hypotheses are also discussed.

Testing the function of petal-carrying in the Red-backed Fairy-wren (Malurus melanocephalus)

In most species of fairy-wren (Malurus spp.), males are known to carry brightly coloured petals and display them to conspecifics. Although petal-carrying by males is often considered an inter-sexual courtship display, anecdotal observations suggest that it may also serve an intra-sexual aggressive function. In this study, we tested hypotheses for the function of petal-carrying displays in the Red-backed Fairy-wren (Malurus melanocephalus) through behavioural observations and aviary-based experiments. All petal displays observed were perpetrated by males, with 90% directed toward females and 10% toward other males. Most petal displays were performed by males in nuptial plumage during intrusions into neighbouring territories, and most displays coincided with the potential fertile period of the female to whom the display was directed. Display items consisted primarily of seeds from Gahnia sp. and flowers from Banksia sp. and Lantana sp., all of which were red or pink and similar to the red backs of nuptial males. Aviary trials found no significant difference in the responses by adult females or adult males to stimulus males with or without petals, substantiating neither a courtship role nor an aggressive role for the display. On the basis of the context of displays in the field, we conclude that the petal-carrying display in the Red-backed Fairy-wren primarily serves an inter-sexual, courtship function, though we could not rule out an intra-sexual, aggressive function.

THE FIRST BASIC PROBLEM: A REVIEW OF MOLT AND PLUMAGE HOMOLOGIES

Abstract All birds have fundamentally similar patterns of plumage succession. Thus Humphrey and Parkes (1959) proposed a system of nomenclature (the H-P system), based on homologies, that has become standard for molt studies in North America. However, presumably analogous similarities in pattern between first basic and definitive basic plumages have obscured homologies. Many plumages conventionally known as “first basic” are better considered as novel first-cycle plumages that lack homologous counterparts in subsequent cycles. Consequently, current nomenclature does not consistently reflect between-species homologies. Howell and Corben (2000b) proposed that traditional juvenal plumage can be considered an unambiguous starting point for a terminology that better reflects presumed homologies in basic plumages; alternate and other nonbasic plumages may not necessarily be homologous between species. Four underlying strategies of increasing complexity incorporate all known patterns of plumage succession: the S...

Costs and benefits of variable breeding plumage in the red-backed fairy-wren.

The red-backed fairy-wren is a socially monogamous passerine bird which exhibits two distinct types of breeding male, bright males that breed in bright red and black plumage and dull males that breed in dull brown plumage. Most males spend their first potential breeding season in dull plumage and subsequent breeding seasons in bright plumage, but a relatively small proportion of males develop bright plumage in their first breeding season. This study quantifies morphology, behavior, and reproductive success of dull and bright males to assess the adaptive costs and benefits of bright plumage while controlling for age. Older bright males (two years of age or older) attempted to increase their reproductive success via copulations with extrapair females, whereas younger (one-year old) bright males and dull males did not. Thus, older bright males spent less time on their own territories, intruded on neighboring groups with fertile females more frequently, gave more courtship displays, and had larger sperm storage organs than did younger bright males and dull males. Microsatellite analyses of paternity indicate that the red-backed fairy-wren has extremely high levels of sexual promiscuity, and that older bright males had higher within-brood paternity than dull males or younger bright males. Regardless of age, bright males were more attractive to females in controlled mate choice trials than were dull males, and both age classes of bright males obtained higher quality mates earlier in the breeding season than did dull males, when nesting success was higher. In conclusion, although it appears that bright plumage increases access to higher quality mates, age also plays a central role in determining a male's overall reproductive success because of the high levels of sexual promiscuity exhibited by the red-backed fairy-wren.

Spring migration strategies of two populations of bar‐tailed godwits, Limosa lapponica, in the Wadden Sea: time minimizers or energy minimizers?

Birds can optimize their migration either by minimizing time of transport, energy expenditure, or predation risk during migration. For each of these optimization criteria different fattening and stopover strategies are predicted. The first two of these optimization criteria are examined here for the bar‐tailed godwit (Limosa lapponica). In the European Wadden Sea two populations of bar‐tailed godwits stop over during spring migration between their wintering and breeding areas. The European population winters mainly in Great Britain and the western part of the Wadden Sea and breeds in Fennoscandia. The Afro‐Siberian population winters in West Africa and breeds in Siberia. The European wintering population migrates to the eastern parts of Wadden Sea in March where it stays until early May. During this time birds gain 1.9 g d−1 in body mass for a 1500–2000‐km non‐stop flight to the breeding areas. Afro‐Siberian birds stay only for one month in May in the Wadden Sea where they gain on average 9.4 g d−1 in mass for a 4000‐km non‐stop flight. Intake rates in April/May did not differ between the two populations (1.5 kJ min−1 and 1.8 kJ min−1 for Siberian and European migrants, respectively) but total energy intake was higher for the Siberian migrants, since they spend 50% of the day foraging vs 30% in the European birds. In contrast to European migrants, Afro‐Siberian birds start to moult into breeding plumage already in their winter quarters. During their stopover in the Wadden Sea thermostatic costs are lower than at times when European birds are present. Thus, the higher energy demands of the Afro‐Siberian birds seem to be fulfilled by a combination of physiological adaptations and a high working level. European birds seem to adopt an energy‐minimized migration strategy whereas Afro‐Siberian birds appear to follow a time‐minimized migration.

Moult, Mass and Flight Range of Waders Ready to Take Off for Long-Distance Migrations

Wader species wintering on the Banc d'Arguin increased their body mass by about 40% during the 4–6 weeks before their departure in spring. This estimate is based on 1) the empirical fact that most waders which had finished or suspended their body moult into summer plumage were heavy and thus ready to take off, and 2) the assumption that only the heaviest birds in the population left, which allowed the mass of disappearing waders to be estimated from counts and the frequency distribution of body masses in samples of captured birds. The mass gain on the Banc d'Arguin is just over 1% per day, when expressed as a proportion of winter mass. A review of studies on waders preparing for migration shows that 1) the total migratory reserve adds 20–80% to winter mass, 2) the rate of mass gain is 0.1–4% per day and 3. the period of mass increase lasts four weeks on average, but longer if waders prepare for spring migration on the wintering areas. We suggest that all wader species leaving the B anc d'Arguin at the end of April and the beginning of May are able to reach SE. and NW. Europe without refuelling. This seems only possible if current equations to predict flight range systematically under-estimate this range, even when the energetic benefits of favourable winds at high altitude are taken into account.

To Leap from Mountains, a Figure of Thought

of course, he said, but it's contingent on physics and chemistry. and provable by rules of mechanics, we tried to plead. he pressed the thumb of his left hand against the pulse of his right, his eyes roaming over the furniture in the room. tinny old pianner, he said, tinny old pianner wrapped in love. although, triggered by my metronome, he said, focusing his eyes through the window on the green of the garden, i used to have a clock in my head and could count on arriving right on the dot, lately i am always late, and it upsets me. it upsets me, he said, as it upsets my search for a new magic of lan guage, forest father of German art, he said, and it reminds me of this childhood game, he said, when we bought a little Chinese doll and squeezed to make it squeak, in very charming contrast to its beautiful clear face with its straight black hair and the delicate colors of its crown of glass beads. it's important to know, he said, and it upsets me. it upsets me that it upsets me, he said. that i'm emotionally dependent on you, quarry and plumage! may his hand rise out of his grave! important to know, he said, and it upsets me. it upsets me, he said, that it upsets me. that i feel persecuted, burdened with persecution by men and things that seem fixated on me so that i in turn begin to be fixated on them, in the hope perhaps to stay clear of them after all, quarry and plum age! what the, he said, bell tolls. just as the tropical moon, he said, which one might liken to a bowl, he said, seemed to pursue me. may his hand rise out of his grave! it upsets me, he said, as it upsets me to leap from mountains in the morning, in hunched position, he said, forever unable to rest, de vouring with mounting impatience all the hours of the day, in fire eating fits all the months of the year. interrupted only by dreams just as breathless, untamed, un bridled, with irregular habits, in hunched position. a trip a trip cut into many small pictures, he said. it upsets me, he said, as it upsets me that i, because alive, try to spew forth everything inside me because i live by living. he felt with the thumb of his right hand the pulse of his left. secondhand happiness, he said, secondhand love. how do you explain this, he said, looking at us and using the formal adress as he did from time to time. czech heart, english hair, he said, how do you explain this. how do you explain this, he said, shoes still covered with dust of new york streets, and already dropped out of the clouds, out of cloudcuckooland, dropped, before brannt. go to hell, he said, go to hell. may his hand rise out of his grave! it upsets me, he said, as it upsets me that i won't be able to go to texas any more, it would have been nice for preserving the texts. …

Social Behavior of the Ruff, Philomachus Pugnax (L.)

INTRODUCTION A. SCOPE OF THE STUDY The Ruff, Philomachus pugnax(L.), differs from other waders in its breeding behavior: courtship and mating take place on a communal display ground or lek; no sounds are uttered during these activities; nesting behavior, incubation, and parental care are the exclusive task of the female. The Ruff is also distinguished by its sexual dimorphism in both size and plumage, and by the extreme individual diversity in the male nuptial plumage. The earliest published accounts of behavior on the lek are those of SELOUS (1906, 1907), a pioneer in the field study of bird behavior. Interested in the Darwinian concept of sexual selection, SELOUS objectively described the interactions among the individual males and females on the lek in an attempt to gather information on partner choice on the lek and, specifically, on the function of figh6ng among the males. His descriptions and interpretations of this behavior are remarkably good considering the date at which they were made. Other accounts of field studies on the breeding behavior of the Ruff have since been published (PORTIELJE 1931; ANDERSEN 1944, 1948, 1951; LINDEMANN 1951; BANCKE & MEESENBURG 1952, 1958; MILDENBERGER 1953), and a number of authors have made comments on different aspects of the Ruff's breeding behavior (e.g., CHRISTOLEIT 1924; ARMSTRONG 1947; TINBERGEN 1959; WYNNE-EDWARDS 1962). VAN OORDT and JUNGE (1934, 1936) have investigated the endocrinological aspects of the development of the male nuptial plumage. LINDEMANN (1951) and GOETHE (1953) describe and comment on the extent of the plumage diversity among the males. The present study extends the observations on the lek behavior and morphological features in the external appearance of the Ruff. Specifically, this is an ethological study of the lek behavior of the Ruff which investigates the social organization of the lek community and the behavior patterns that serve mutual communication among the birds. It is assumed that the behavior patterns provide many clues for the understanding of the social organization. Further, particular attention is paid to the biological significance of the plumage diversity within the lek community. Accidentally, this study revealed that the Ruff presents an extreme example of behavioral polymorphism: it appears that within the male community two groups of males coexist which show distinct differences in behavior. In a complicated way these behavioral differences are associated with differences in the external appearance of the individual males. This paper is organized in the following manner. Descriptive data on morphologicai features in the external appearance of the Ruff are presented in Chapter II. Mutual relations within the male community, correlations of behavioral differences with external appearance, and the intersexual relations during courtship are dealt with in Chapter III; the typical situations on the lek in which the behavior patterns occur are also described in this chapter. Detailed descriptions of the form of these behavior patterns are presented in Chapter IV. Chapter V is concerned with a discussion of different aspects of the behavioral polymorphism and the plumage diversity in the male Ruff. ... Zie: Introduction

UNUSUAL SUMMER PLUMAGE OF WHITE-TAILED PTARMIGAN

We observed two White-tailed Ptarmigan (Lagopus leucurus) in white winter plumage (basic) in Colorado during late July through August during 35 years of field study. Both individuals were adult (>2 yr of age) females; one was collected and the second examined and photographed. Examination of the plumage revealed that neither had molted from winter plumage into their nuptial (alternate) or postnuptial (supplemental) plumage. We speculate that retention of the white winter plumage was the result of failure of the pituitary and thyroid glands to secrete sufficient hormones to initiate replacement of white body feathers with normal breeding plumage feathers. Perhaps this failure of the pituitary and thyroid glands was not age related but instead gender related. Despite conspicuous white plumage during summer, behavior and survival of both birds appeared normal.