Papers by Theunis Piersma
Journal of Ornithology, 1994
Journal of Ornithology, 1994
: Unraveling the intraspecific phylogeography of Knots Calidris canutus: a progress report on the... more : Unraveling the intraspecific phylogeography of Knots Calidris canutus: a progress report on the search for genetic markers. J. Orn. 135: 599-608. -Mitochondrial DNA control region sequences of 25 Knots sampled from 10 populations and possibly four subspecies (canutus, islandica, rogersi, tufa) were obtained by PCR and direct sequencing. OnIy 7 haplotypes were found worldwide, all closely related to one another and differing by 1-3 substitutions. Knots have most probably expanded to their current population size from a refugial population that was severely bottlenecked late in the Pleistocene. Preliminary results from RAPDs are consistent with this prediction, in that Knots from North America appear to be genetically distinct from Knots elsewhere.
Journal of Ornithology, 1986

Journal of Ornithology, 2011
We isolated and tested six novel microsatellite loci in Sanderling (Calidris alba) from Greenland... more We isolated and tested six novel microsatellite loci in Sanderling (Calidris alba) from Greenland for paternity analyses. In addition, we tested 11 already published microsatellite markers which were originally developed for the congeneric species, the Pectoral Sandpiper (C. melanotos). All loci were polymorphic, but five of the cross-species loci were not scorable due to suboptimal amplification patterns. The 12 successful loci were tested on 87 individuals, yielding an average of 9.0 (range 4-19) alleles per locus and mean expected heterozygosity of 0.70. Because this dataset contained families, tests for Hardy-Weinberg equilibrium, linkage disequilibrium and probability of identity were done on a subset of the data containing 25 adults caught in the same year. The overall probability of identity was 1.0 9 10 -13 . Only one locus displayed significant homozygote excess and all loci were unlinked. On the basis of female heterozygotes, all loci are assumed to be autosomal.
Journal of Ornithology, 2006

Journal of Ornithology, 2007
Shorebirds, or waders, form an ecologically (but not phylogenetically) homogenous group of birds ... more Shorebirds, or waders, form an ecologically (but not phylogenetically) homogenous group of birds that, despite this homogeneity, exhibits clear correlated contrasts in habitat use and migration distance between closely related species pairs. In addition, within species there is distinct variation in breeding and wintering latitudes, i.e. migration distance. I examine here such contrasts at different taxonomic levels and evaluate what we can learn about selective forces on habitat selection and the evolution of migration strategies in birds. My primary example is the worldwide migration system of the Red Knot Calidris canutus. These sandpipers breed only on high arctic tundra (65-83°N), but they move south from their disjunct, circumpolar breeding areas to nonbreeding sites on the coasts of all continents (except Antarctica), between latitudes 58°N and 53°S. Due to their specialized sensory capabilities, Red Knots generally eat hard-shelled prey found on intertidal, mostly soft, substrates. As a consequence, ecologically suitable coastal sites are few and far between, so they must routinely undertake flights of many thousands of kilometres. In contrast to prediction, Red Knots at tropical intertidal sites have lower fuelling rates than birds at more southern or northern latitudes. This leads to greater time-stress in the southernmost wintering populations that not only have to cover over 14,000 km in single migrations, but also cannot rely on tropical regions to make refuelling stops. Rapid human-caused losses of the food-base in staging areas during both north-and southward migrations have been demonstrated to have caused rapid declines in several Red Knot populations. Detailed worldwide ecodemographic research on these extreme long-distance migrants, as embodied in, for example, the recently established Global Flyway Network, yields a two-pronged benefit: (1) on the basis of the unintended large-scale experiments carried out by humans, we rapidly come to grips with the selection pressures moulding the migration strategies of migrant birds, and (2) in applied contexts, the work gives instantaneous feedbacks on the conservation consequences of man-made alterations to wetland environments at the relevant global spatial scales.

Journal of Ornithology, 2011
When early breeding is advantageous, migrants underway to the breeding areas may be time stressed... more When early breeding is advantageous, migrants underway to the breeding areas may be time stressed. The timing of sequential events such as migration and breeding is expected to be correlated because of a ''domino effect'', and would be of particular biological importance if timings are repeatable within individuals between years. We studied a colour-marked population of Black-tailed Godwits Limosa l. limosa both on staging areas in Portugal and on breeding areas in The Netherlands. For each individual, we measured the timing of the staging period, the arrival date on the breeding area and the egg laying date. We measured average egg volume as a measure of reproductive investment. The date of departure from the staging areas, the arrival date on the breeding areas, and the egg laying date were repeatable among years in individual Black-tailed Godwits. The arrival dates of paired males and females and the average annual male and female arrival dates were correlated. The dates of departure from Portugal, arrival in The Netherlands, and egg laying were not correlated. Earlier clutches had larger eggs than late clutches. If the length of the individually available pre-laying period is accounted for, early arriving birds spent more time on the breeding grounds before laying than late arriving birds. The repeatability of the itineraries and the correlation between arrival timing of males and females are consistent with observations in other migrants. Despite evidence for early breeding being advantageous, we found no evidence of a ''domino effect''.
Journal of Natural Products, 2000
The intact C 32 -C 48 diester wax esters of the preen gland of the migrating bird Calidris canutu... more The intact C 32 -C 48 diester wax esters of the preen gland of the migrating bird Calidris canutus are shown, using synthesized standards, to comprise predominantly C 12 -C 16 alkane-1,2-diols esterified with octanoic, decanoic, and dodecanoic acid at one position, and with predominantly even-numbered carbon fatty acids at the other position.

Journal of Experimental Zoology Part A: Ecological Genetics and Physiology, 2007
Birds living in seasonal environments change physiology and behavior in correspondence to tempora... more Birds living in seasonal environments change physiology and behavior in correspondence to temporally changing environmental supplies, demands and opportunities. We recently reported that the chemical composition of uropygial gland secretions of sandpipers (Scolopacidae, order Charadriformes) changes during the breeding season from mixtures of monoesters to diesters, which fulfill specific functions related to incubation. A proper temporal match between the expression of diester preen waxes and incubation requires a flexible organization of the trait. Here we analyze the possible degrees of flexibility with reference to the functionality of better-understood molt and body mass cycles of free-living and captive red knots (Calidris canutus). The relative flexibility of seasonal cycles in preen wax composition was examined by two experimental perturbations: (1) giving birds restricted access to food and (2) monitoring them long-term under a constant photoperiodic regime. We found that wax type cannot change instantaneously, but that changing the type of wax is under similar organizational time constraints as the replacement of feathers. Just as molt and mass cycles, the seasonal rhythm of diester secretion appeared to be under endogenous control: most birds placed in a constant photoperiod still maintained seasonally changing preen waxes. Diester preen wax secretion was synchronized with the peak in body mass in spring, but became less well expressed under constant photoperiodic conditions and when food availability was limited.
Journal of Experimental Marine Biology and Ecology, 2011
Daily growth rate in situ fluorescent marking Intertidal mudflat Lucinidae Mark-and-recapture Sea... more Daily growth rate in situ fluorescent marking Intertidal mudflat Lucinidae Mark-and-recapture Seagrass

Journal of Experimental Marine Biology and Ecology, 2008
In marine bivalves, the relative sizes of the gills and palps appear to be a useful functional tr... more In marine bivalves, the relative sizes of the gills and palps appear to be a useful functional trait that reflect feeding mode, i.e. suspension feeders have relatively larger gills than palps for pumping, whereas deposit feeders have relatively larger palps than gills for sorting. Also, within a species, the relative sizes of the gills and palps are related to changes in local food conditions. However, there is still no firm evidence showing that differences in the relative gill and palp sizes between species are related to diet selection. Based on the knowledge that carbon and nitrogen isotope signatures of an animals tissues reflect past diet, we compared the relative gill and palp sizes of bivalves from Roebuck Bay, northwestern Australia with their carbon and nitrogen isotope signatures. The carbon isotope signatures distinguished clear differences in diet between bivalves along a gradient from suspension to deposit feeding, and strikingly this pattern was closely followed by the relative sizes of the gills and palps of the bivalves. This study confirms that relative gill and palp sizes in bivalves are a functional trait that can be used to compare resource use between species. Furthermore, these data may suggest that morphospace occupation, as determined by relative gill and palp sizes of bivalves, could reflect a gradient of resource use between species.

Journal of Experimental Marine Biology and Ecology, 2007
The climate variability hypothesis proposes that in variable temperate climates poikilothermic an... more The climate variability hypothesis proposes that in variable temperate climates poikilothermic animals have wide thermal tolerance windows, whereas in constant tropical climates they have small thermal tolerance windows. In this study we quantified and compared the upper and lower lethal thermal tolerance limits of numerous bivalve species from a tropical (Roebuck Bay, north western Australia) and a temperate (Wadden Sea, north western Europe) tidal flat. Species from tropical Roebuck Bay had higher upper and lower lethal thermal limits than species from the temperate Wadden Sea, and Wadden Sea species showed an ability to survive freezing temperatures. The increased freezing resistance of the Wadden Sea species resulted in thermal tolerance windows that were on average 7°C greater than the Roebuck Bay species. Furthermore, at a local-scale, the upper lethal thermal limits of the Wadden Sea species were positively related to submersion time and thus to encountered temperature variation, but this was not the case for the Roebuck Bay species. A review of previous studies, at a global scale, showed that upper lethal thermal limits of tropical species are closer to maximum habitat temperatures than the upper lethal thermal limits of temperate species, suggesting that temperate species are better adapted to temperature variation. In this study, we show for the first time, at both local and global scales, that the lethal thermal limits of bivalves support the climate variability effect in the marine environment.

Journal of Evolutionary Biology, 2003
Understanding the origin, maintenance and significance of phenotypic variation is one of the cent... more Understanding the origin, maintenance and significance of phenotypic variation is one of the central issues in evolutionary biology. An ongoing discussion focuses on the relative roles of isolation and selection as being at the heart of genetically based spatial variation. We address this issue in a representative of a taxon group in which isolation is unlikely: a marine broadcast spawning invertebrate. During the free-swimming larval phase, dispersal is potentially very large. For such taxa, small-scale population genetic structuring in neutral molecular markers tends to be limited, conform expectations. Small-scale differentiation of selective traits is expected to be hindered by the putatively high gene flow. We determined the geographical distribution of molecular markers and of variation in a shell shape measure, globosity, for the bivalve Macoma balthica (L.) in the western Dutch Wadden Sea and adjacent North Sea in three subsequent years, and found that shells of this clam are more globose in the Wadden Sea. By rearing clams in a common garden in the laboratory starting from the gamete phase, we show that the ecotypes are genetically different; heritability is estimated at 23%. The proportion of total genetic variation that is between sites is much larger for the morphological additive genetic variation (Q ST ¼ 0.416) than for allozyme (F ST ¼ 0.000-0.022) and mitochondrial DNA cytochrome-c-oxidase-1 sequence variation (U ST ¼ 0.017). Divergent selection must be involved and intraspecific spatial genetic differentiation in marine broadcast spawners is apparently not constrained by low levels of isolation.

Journal of Comparative Physiology B, 1995
Seasonality in basal metabolic rate and thermal conductance in a long.distance migrant shorebird,... more Seasonality in basal metabolic rate and thermal conductance in a long.distance migrant shorebird, the knot (Calidris canutus) Abstract Knots Calidris canutus live highly seasonal lives, breeding solitarily on high arctic tundra and spending the non-breeding season in large social flocks in temperate to tropical estuaries. Their reproductive activities and physiological preparations for long flights are reflected in pronounced plumage and body mass changes, even in long-term captives of the islandica subspecies (breeding in north Greenland and northeast Canada and wintering in western Europe) studied in outdoor aviaries. The three to four fattening episodes in April-July in connection with the flights to and from the high arctic breeding grounds by freeliving birds, are represented by a single period of high body mass, peaking between late May and early July in a sample of ten captive islandica knots studied over four years. There are consistent and synchronized annual variations in basal metabolic rate and thermal conductance in three istandica knots. Basal metabolic rate was highest during the summer body mass peak. Within the examined individuals, basal metabolic rate scales on body mass with an exponent of about 1.4, probably reflecting a general hypertrophy of metabolically expensive muscles and organs. Any potential effect of moult on basal metabolic rate was obscured by the large seasonal mass-associated variations. In breeding plumage, insulation (the inverse of thermal conductance) was a factor of 1.35 lower than in winter plumage. This was paralleled by the dry mass of contour T. Piersma (~) . N. feathers being a factor of 1.17 lower. In this subspecies the breeding season is indeed the period during which the costs of thermoregulation are lowest. In captive knots seasonal changes in basal metabolic rate and thermal conductance likely reflect an anticipatory programme adaptive to the variable demands made by the environment at different times of the year.

Journal of Comparative Physiology A, 2009
The winter immunoenhancement hypothesis associates long nights and increased exposure to melatoni... more The winter immunoenhancement hypothesis associates long nights and increased exposure to melatonin with enhanced immune function in winter when resource availability is low and the chances of becoming ill are high. Thus, increased exposure to melatonin in the winter could be adaptive for species facing diYcult winter conditions. This idea has found some support in studies of resident mammals. In birds, the link between day length and melatonin over the annual cycle is weaker, and contributions of melatonin to seasonal timing are unclear. Furthermore, many species, especially migrants, do not experience the most diYcult conditions of their annual cycle in winter. In this study, we tested whether the winter immunoenhancement hypothesis holds in an avian species, the red knot Calidris canutus. We found that melatonin duration and amplitude varied signiWcantly over the annual cycle with the highest values occurring in winter. However, peaks did not correspond to the winter solstice or with annual variation in immune function. Our Wndings do not support the winter immunoenhancement hypothesis in knots and question whether the idea that immune function should be bolstered in winter can be generalized to systems where winter is not the most diYcult time of the year.

Journal of Avian Biology, 2002
Bar-tailed godwits migrate from West African wintering sites to breeding areas in northern Russia... more Bar-tailed godwits migrate from West African wintering sites to breeding areas in northern Russia with only one stopover. We compared hematocrit (Hct), blood hemoglobin concentration (Hb), and mean cell hemoglobin concentration (MCHb; a measure of the relative proportion of Hb in the cellular blood fraction) between arriving godwits lured to land 60 km short of the stopover site and godwits during subsequent refueling. The Hct and Hb of arriving godwits was low when compared to that of refueling birds. On the stopover site, Hct and Hb correlated positively with size-corrected body mass. In addition, Hb and MCHb reached peak levels in the last days of stopover. We explored the possibility of regenerative anemia in arriving godwits by comparing the fraction of reticulocytes (young red blood cells) between arriving and refueling birds. No differences were found. Therefore, we suggest that the increase in Hct, Hb, and MCHb during refueling is not in response to a severe anemic state at arrival. Rather, we suggest that the increase in blood parameters may anticipate the increased aerobic requirements of impending migratory flight and possibly satisfy heightened oxygen demands of the larger body mass of fattened birds. The Hct increase on the stopover site may also serve to buffer the red blood cell population against possible red blood cell breakdown during long-distance flight.
Ruddy turnstones Arenaria interpres rapidly build pectoral muscle after raptor scares
Journal of Avian Biology, 2006
... 2). A threat of predation was induced by gliding the raptor along a line along one wall of th... more ... 2). A threat of predation was induced by gliding the raptor along a line along one wall of the aviary, once a day but at ... Calculated pectoral-free lean mass changed from 52.6 g at the start of trials, to 50.7 g after raptor scares and 51.3 g after the ... John Wiley & Sons, New York. ...
Journal of Avian Biology, 2005
Phenotypic Flexibility during Migration: Optimization of Organ Size Contingent on the Risks and Rewards of Fueling and Flight?
Journal of Avian Biology, 1998
... Using preliminary comparative data for different populations of Bar-tailed Godwits Limosa lap... more ... Using preliminary comparative data for different populations of Bar-tailed Godwits Limosa lapponica, and several other shorebird species adding variation to the migration strategies sampled (Golden Plover Pluvialis apricaria, Ruff Philomachus pugnax, Red Knot Calidris ...
Journal of Avian Biology, 1996
JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, a... more JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. . Wiley and Nordic Society Oikos are collaborating with JSTOR to digitize, preserve and extend access to Journal of Avian Biology. Weber, T. P. and Piersma, T. 1996. Basal metabolic rate and the mass of tissues differing in metabolic scope: migration-related covariation between individual Knots Calidris canutus. -J. Avian Biol. 27: 215-224.
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Papers by Theunis Piersma