Continuous, age-related plumage variation in male Kirtland's Warblers

ABSTRACT.   The ability to age individual birds visually in the field based on plumage variation could provide important demographic and biogeographical information. We describe an approach to infer ages from a distribution of plumage scores of free-ranging male Kirtland's Warblers ( Dendroica kirtlandii ). We assigned ages to males using a scoring scheme (0–12 points) based on variation in plumage coloration, brightness, and contrast on three dorsal and three ventral body regions presumed to be age-related. The distribution of total additive plumage scores for 875 breeding males was normally distributed, indicating no distinct age classes. Thus, we developed provisional plumage-age classes of second year (SY) and after second-year (ASY), and compared them to the total plumage scores of a smaller subsample of known age ( N = 92) and minimum age ( N = 143) males. Plumage scores of known-age male Kirtland's Warblers increased nonlinearly with age ( r _s= 0.67), but with some overlap. The median plumage score for SY males (median = 5.0) was significantly lower than for third-year (TY) males (median = 7.0) and after third-year (3 year and older) males (median = 8.0), indicating that the plumage of male Kirtland's Warblers becomes more distinctive and brighter with age. Linear discriminant function analysis differentiated ASY male Kirtland's Warbler from SY males with 78.3% accuracy. Investigators could use the distribution of plumage scores and approximate age structures to document changes in male age structure during colonization, use, and abandonment of habitats by Kirtland's Warblers or other species that occupy early successional habitats. Aging free-ranging birds based on a plumage scoring scheme may be especially critical for demographic studies of less-studied species where it is unlikely that a banding program will be initiated, but where plumage-age inferences or management decisions must be made.     

Rapid recovery of Dutch gray seal colonies fueled by immigration

Gray seals were first observed breeding in the Dutch Wadden Sea in 1985, after centuries of absence. The breeding colony there is now the largest on the European continent. We describe the changes in gray seal numbers and their geographical expansion, and estimate how these processes were influenced by immigration from other colonies. Counts of hauled out animals were carried out between 1985 and 2013, monitoring three different periods of the seals' annual cycle. Using priors determined for the UK population, a Bayesian demographic model was fitted to pup numbers to estimate the population parameters driving the growth. This included immigration of subadults into the breeding population, which contributed to an average growth rate in the pup counts of 19%/yr, much higher than expected in a closed population. This immigration may account for approximately 35% of the total annual growth. In addition, at least 200 gray seals from the UK visit the area temporarily. Recovery of the population in the Netherlands occurred more than 50 yr after gray seals were protected in the UK. These time scales should be taken into account when studying long living marine mammals, e.g., in impact and conservation studies.     

Postembryonic development in Diaphanosoma brachyurum (Lievin, 1848) (Crustacea: Ctenopoda: Sididae)

Postembryonic females and males Diaphanosoma brachyurum from Lake Glubokoe (Moscow) have 3–4 and 3 juvenile instars, respectively. Females and males of the first three postembryonic instars can be identified by the different number of setae and setal rudiments on the proximal and distal segments of the exopodite of the swimming antennae: 3 + 7; (i + 3) + 7; 4 + (i + 7), respectively (i = rudiment of seta). The subsequent instars have 4 + 8 long plumose setae on these segments, but the fourth instar has the proximal lateral seta of the distal exopod segment slightly shorter and thinner than the others. The antennules and copulatory appendages of males are instar-specific. Diaphanosomas show small increments in body length during the postembryonic molts. The largest increments (about 115 m) occur during the first or second molts. The allometric equation of Huxley (1924) was used for a comparison of the relative growth rate of different body parts. In the middle of summer, the head and swimming antennae with the body and the antennal exopodite with the antennal basipodite grow in isometry. At the same time, the branches of the swimming antennae and their setae show allometric growth: the exopodite and distal setae grow faster than the endopodite and the lateral setae, respectively.     

Flexibility and constraints in the molt schedule of long‐distance migratory shorebirds: causes and consequences

Molt is a major component of the annual cycle of birds, the timing and extent of which can affect body condition, survival, and future reproductive success through carry‐over effects. The way in which molt is fitted into the annual cycle seems to be a somewhat neglected area which is both of interest and of importance. Study of the causes of annual variation in the timing of molt and its potential consequence in long‐distance migratory birds was examined using the Curlew Sandpiper, Calidris ferruginea, as a model species. Using the maximum likelihood molt models of Underhill and Zucchini (1988, Ibis 130:358–372), the relationship between annual variability in the start dates of molt at the population level with conditions on the breeding area was explored. Adult males typically started early in years when temperature in June on the Arctic breeding grounds were high compared to cold years while adult females molted later in years of high breeding success and/or warm July temperature and vice versa. When molt started later, the duration was often shorter, indicating that late completion of molt might have fitness consequences, probably jeopardizing survival. Evidence of this was seen in the low body condition of birds in years when molt was completed late. The results indicate that these migratory shorebirds follow a fine‐tuned annual life cycle, and disturbances at a certain stage can alter next biological events through carry‐over effects.     

Complex postbreeding molt strategies in a songbird migrating along the East Asian Flyway,the Pallas’s Grasshopper Warbler Locustella certhiola

Molt strategies have received relatively little attention in current ornithology, and knowledge concerning the evolution, variability and extent of molt is sparse in many bird species. This is especially true for East Asian Locustella species where assumptions on molt patterns are based on incomplete information. We provide evidence indicating a complex postbreeding molt strategy and variable molt extent among the Pallas's Grasshopper Warbler Locustella certhiola, based on data from six ringing sites situated along its flyway from the breeding grounds to the wintering areas. Detailed study revealed for the first time that in most individuals wing feather molt proceeds from the center both toward the body and the wing‐tip, a molt pattern known as divergent molt (which is rare among Palearctic passerines). In the Russian Far East, where both breeding birds and passage migrants occur, a third of the adult birds were molting in late summer. In Central Siberia, at the northwestern limit of its distribution, adult individuals commenced their primary molt partly divergently and partly with unknown sequence. During migration in Mongolia, only descendantly (i.e., from the body toward the wing‐tip) molting birds were observed, while further south in Korea, Hong Kong, and Thailand the proportion of potential eccentric and divergent feather renewal was not identifiable since the renewed feathers were already fully grown as expected. We found an increase in the mean number of molted primaries during the progress of the autumn migration. Moderate body mass levels and low‐fat and muscle scores were observed in molting adult birds, without any remarkable increase in the later season. According to optimality models, we suggest that an extremely short season of high food abundance in tall grass habitats and a largely overland route allow autumn migration with low fuel loads combined with molt migration in at least a part of the population. This study highlights the importance of further studying molt strategy as well as stopover behavior decisions and the trade‐offs among migratory birds that are now facing a panoply of anthropogenic threats along their flyways.     

The dynamics of exoskeletal-epidermal structure during molt in juvenile lobster by electron microscopy and electron spectroscopic imaging

The exoskeletal-epidermal complex of juvenile lobsters at various stages throughout the molt cycle was examined by conventional electron microscopy, freeze-etch replicas, and electron spectroscopic imaging. This latter technique which enables the direct localization of atomic elements superimposed over morphological fine structure has been applied to this tissue complex to determine the spatial distributions and interrelationships of calcium, phosphorus, and sulphur. Chitin microfibril assembly is visualized in thin sections as occurring at the surface of apical membrane plaques which in freeze-etch replicas invariably possess a rich distribution of intramembrane particles on both P and E faces. In early stages of mineralization the exo- and endocuticular zones of the exoskeleton possess a dense Ca-containing lamellar repeat. These bands are unrelated to the helicoidal arrangement of chitin microfibrils. At later stages of development mineral deposits occur within the exocuticle and advance through to the endocuticle. These deposits align with chitin microfibrils and exhibit a helicoidal pattern. Morphological and chemical alterations associated with mineralization and demineralization of the exoskeleton are discussed.     

Interactive effects of rutin and constant versus alternating temperatures on performance ofManduca sexta caterpillars

The effects of different concentrations of rutin and constant temperature (20 °C) versus alternating temperatures (23∶15 °C) on growth, molting and food utilization efficiencies of third instar tobacco hornworms (Manduca sexta) were determined. Relative consumption rate (RCR) and relative growth rate (RGR) were significantly higher for larvae at the alternating thermal regime compared to those at the constant (representing the average) temperature. With increasing concentrations of rutin, the negative effect of rutin on RCR and RGR increased for the larvae in the alternating thermal regime; however, at the constant temperature, rutin had little effect. The alternating thermal regime promoted synchrony in the timing of spiracle apolysis (the earliest morphological marker of molt). Rutin disrupted that synchrony. I discuss how patterns of host plant resistance may be altered with a decrease, in amplitude of diurnal temperatures (as has been documented recently for temperate regions) through the uncoupling of herbivore performance and allelochemical concentration. I conclude that simultaneous consideration of fluctuating temperatures and allelochemicals is advisable when assessing the effects of temperature and allelochemicals on performance of insect herbivores because interactive effects between temperature and dietary components occur and perhaps are common.     

Variation in foraging investment during the intermolt interval and before egg-laying in the spiderNephila clavipes (Araneae: Araneidae)

Repeated observations were made of marked, freeliving females in three disjunct populations of the orb-weaving spider Nephila clavipes,recording orb size and structure and barrier web complexity. These longitudinal studies revealed regular patterns of variation in orb size associated with the molting cycle and cycle of egg maturation. Juveniles maintained in an insectary under a controlled feeding regime exhibited similar variation in orb size. All spiders ceased orb renewal prior to molting and the majority replaced the viscid orb with a nonviscid platform prior to molting. Differences among the populations in molting web structure are discussed in light of differences in the physical and biological environments. Cessation of orb renewal corresponded temporally with reported physiological changes associated with the molt cycle and possibly reflect changes in resource partitioning between growth and foraging during the molt cycle.     

Membrane Particle Distribution in the Sternal Epithelia of the Terrestrial Isopod Porcellio scaber Latr. (Crustacea, Oniscidea) during CaCO3 Deposit Formation and Resorption, a Freeze-Etch Analysis

The anterior sternal epithelium of terrestrial isopods transports cuticular Ca²⁺ to and from large sternal CaCO₃ deposits. We analyzed the anterior and posterior sternal epithelium by the means of the freeze-etch technique and measured the size distribution and density of intramembrane particles (IMPs) during three different molting stages. At least three IMP size classes around 4.5, 7.7, and 9.4 nm can be distinguished on the P-face of the apical and basolateral plasma membrane. An additional size class of around 12.8 nm is restricted to the apical compartment. In the anterior sternal epithelium, the density of these large particles changes by a factor of 1.9 during the molt cycle, suggesting a role in CaCO₃ formation and/or resorption. The density of the smaller IMPs rises transiently by a factor of 1.3 in the posterior sternal epithelium only. The IMP density of the basolateral plasma membrane increases significantly by a factor of 1.4 and 1.3 in the anterior and posterior sternal epithelia, respectively. The results indicate that increases in the IMP density contribute to the differentiation to an increased transport activity during the cyclic enlargements of the plasma membrane surface area in the anterior sternal epithelium.     

GROWTH RATE AND SHEDDING OF VIBRISSAE IN THE GRAY SEAL, HALICHOERUS GRYPUS: A CAUTIONARY NOTE FOR STABLE ISOTOPE DIET ANALYSIS

Stable isotopes have become powerful tools for gathering information on food webs in marine ecosystems. The method is based on the concept that the ratio of Nitrogen-14 to ¹⁵N (or Carbon-12 to ¹³C) in the tissues of animals is directly related to the ratio found in their diet. Vibrissae provide a time series of stable isotope data as tissue is laid down sequentially over time. Here we examine the growth rate of 283 mystacial (muzzle) vibrissae of four gray seals, Halichoeruas grypus , over a five-month period to investigate their applicability for stable isotope diet analysis. The individual vibrissae did not grow at a constant rate during the study, Fifty-nine actively growing vibrissae were modeled to quantify the growth pattern using a three-parameter von Bertalanffy curve, with the parameters estimated using non-linear mixed-effects models. This model incorporated the inherent serial correlation of these data. The growth rate was 0.024 cm/d (95% CI = 0.019–0.030), the asymptotic length differed significantly by location ( F _3,56=9.64, P < 0.001), but no significant trend was found with muzzle location ( F _3,56= 0.15, P = 0.93). The Δlength/Δtime between each measurement was calculated and most of these data fell at or near zero growth (median = 0.04 cm/d, range = 0–0.78). Individual vibrissae were shed asynchronously and without any seasonal growth trend. This has serious implications for researchers attempting to extrapolate diet data from vibrissae. Because the growth is neither continuous nor synchronous, it will be a challenge to accurately identify the dates when the isotopes were incorporated into the tissue.