The relationship between limb morphology, kinematics, and force during running: the evolution of locomotor dynamics in lizards

Terrestrial locomotion occurs via the hierarchical links between morphology, kinematics, force, and center-of-mass mechanics. In a phylogenetically broad sample of seven lizard species, we show that morphological variation drives kinematic variation, which, in turn, drives force variation. Species with short limbs use a short stride–high frequency strategy when running at steady-speed and to change speeds. This link between morphology and kinematics results in relatively small vertical forces during the support phase of the stride cycle. Conversely, species with long limbs use a long stride–low frequency strategy, resulting in large vertical forces during the support phase. In view of these findings, we suggest that limb length may predict locomotor energetics in lizards because energetics are largely determined by vertical forces and stride frequency. Additionally, we propose an energetic trade-off with both long- and short-limbed species paying the most energy to move, whereas intermediate-limbed species move using less energy. Finally, when these traits are mapped onto a lizard phylogeny, we show that locomotor functional morphology exhibits both deep phylogenetic effects and contemporary patterns of evolutionary convergence. Overall, the present study provides a foundation for testing hypotheses regarding the integration and evolution of functional traits in lizards and animals in general.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 634–651.     

Satellite-marked waterfowl reveal migratory connection between H5N1 outbreak areas in China and Mongolia

The role of wild birds in the spread of highly pathogenic avian influenza H5N1 has been greatly debated and remains an unresolved question. However, analyses to determine involvement of wild birds have been hindered by the lack of basic information on their movements in central Asia. Thus, we initiated a programme to document migrations of waterfowl in Asian flyways to inform hypotheses of H5N1 transmission. As part of this work, we studied migration of waterfowl from Qinghai Lake, China, site of the 2005 H5N1 outbreak in wild birds. We examined the null hypothesis that no direct migratory connection existed between Qinghai Lake and H5N1 outbreak areas in central Mongolia, as suggested by some H5N1 phylogeny studies. We captured individuals in 2007 from two of the species that died in the Qinghai Lake outbreaks and marked them with GPS satellite transmitters: Bar-headed Geese Anser indicus ( n  =   14) and Ruddy Shelduck Tadorna ferruginea ( n  =   11). Three of 25 marked birds (one Goose and two Shelducks) migrated to breeding grounds near H5N1 outbreak areas in Mongolia. Our results describe a previously unknown migratory link between the two regions and offer new critical information on migratory movements in the region.     

Acclimation of leaf hydraulic conductance and stomatal conductance of Pinus taeda (loblolly pine) to long-term growth in elevated CO2 (free-air CO2 enrichment) and N-fertilization

We investigated how leaf hydraulic conductance (K_leaf) of loblolly pine trees is influenced by soil nitrogen amendment (N) in stands subjected to ambient or elevated CO₂ concentrations (CO₂^a and CO₂^e, respectively). We also examined how K_leaf varies with changes in reference leaf water potential (Ψ_leaf‐ref) and stomatal conductance (g_s‐ref) calculated at vapour pressure deficit, D of 1 kPa. We detected significant reductions in K_leaf caused by N and CO₂^e, but neither treatment affected pre‐dawn or midday Ψ_leaf. We also detected a significant CO₂^e‐induced reduction in g_s‐ref and Ψ_leaf‐ref. Among treatments, the sensitivity of K_leaf to Ψ_leaf was directly related to a reference K_leaf (K_leaf‐ref computed at Ψ_leaf‐ref). This liquid‐phase response was reflected in a similar gas‐phase response, with g_s sensitivity to D proportional to g_s‐ref. Because leaves represented a substantial component of the whole‐tree conductance, reduction in K_leaf under CO₂^e affected whole‐tree water use by inducing a decline in g_s‐ref. The consequences of the acclimation of leaves to the treatments were: (1) trees growing under CO₂^e controlled morning leaf water status less than CO₂^a trees resulting in a higher diurnal loss of K_leaf; (2) the effect of CO₂^e on g_s‐ref was manifested only during times of high soil moisture.     

Identification and functional characterization of effectors in expressed sequence tags from various life cycle stages of the potato cyst nematode Globodera pallida

In this article, we describe the analysis of over 9000 expressed sequence tags (ESTs) from cDNA libraries obtained from various life cycle stages of Globodera pallida . We have identified over 50 G. pallida effectors from this dataset using bioinformatics analysis, by screening clones in order to identify secreted proteins up-regulated after the onset of parasitism and using in situ hybridization to confirm the expression in pharyngeal gland cells. A substantial gene family encoding G. pallida SPRYSEC proteins has been identified. The expression of these genes is restricted to the dorsal pharyngeal gland cell. Different members of the SPRYSEC family of proteins from G. pallida show different subcellular localization patterns in plants, with some localized to the cytoplasm and others to the nucleus and nucleolus. Differences in subcellular localization may reflect diverse functional roles for each individual protein or, more likely, variety in the compartmentalization of plant proteins targeted by the nematode. Our data are therefore consistent with the suggestion that the SPRYSEC proteins suppress host defences, as suggested previously, and that they achieve this through interaction with a range of host targets.     

Population genetic analysis of a recent range expansion: mechanisms regulating the poleward range limit in the volcano barnacle Tetraclita rubescens

As range shifts coincident with climate change have become increasingly well documented, efforts to describe the causes of range boundaries have increased. Three mechanisms—genetic impoverishment, migration load, or a physical barrier to dispersal—are well described theoretically, but the data needed to distinguish among them have rarely been collected. We describe the distribution, abundance, genetic variation, and environment of Tetraclita rubescens, an intertidal barnacle that expanded its northern range limit by several hundreds of kilometres from San Francisco, CA, USA, since the 1970s. We compare geographic variation in abundance with abiotic and biotic patterns, including sea surface temperatures and the distributions of 387 co‐occurring species, and describe genetic variation in cytochrome c oxidase subunit I, mitochondrial noncoding region, and nine microsatellite loci from 27 locations between Bahia Magdalena (California Baja Sur, Mexico) and Cape Mendocino (CA, USA). We find very high gene flow, high genetic diversity, and a gradient in physical environmental variation coincident with the range limit. We infer that the primary cause of the northern range boundary in T. rubescens is migration load arising from flow of maladapted alleles into peripheral locations and that environmental change, which could have reduced selection against genotypes immigrating into the newly colonized portion of the range, is the most likely cause of the observed range expansion. Because environmental change could similarly affect all taxa in a region whose distributional limits are established by migration load, these mechanisms may be common causes of range boundaries and largely synchronous multi‐species range expansions.     

Fat frogs,mobile genes: unexpected phylogeographic patterns for the ornate chorus frog (Pseudacris ornata)

The southeastern coastal plain of the United States is a region marked by extraordinary phylogeographic congruence that is frequently attributed to the changing sea levels that occurred during the glacial‐interglacial cycles of the Pleistocene epoch. A phylogeographic break corresponding to the Apalachicola River has been suggested for many species studied to date that are endemic to this region. Here, we used this pattern of phylogeographic congruence to develop and test explicit hypotheses about the genetic structure in the ornate chorus frog (Pseudacris ornata). Using 1299 bp of mtDNA sequence and seven nuclear microsatellite markers in 13 natural populations of P. ornata, we found three clades corresponding to geographically distinct regions; one spans the Apalachicola River (Southern Clade), one encompasses Georgia and South Carolina (Central Clade) and a third comprises more northerly individuals (Northern Clade). However, it does not appear that typical phylogeographic barriers demarcate these clades. Instead, isolation by distance across the range of the entire species explained the pattern of genetic variation that we observed. We propose that P. ornata was historically widespread in the southeastern United States, and that a balance between genetic drift and migration was the root of the genetic divergence among populations. Additionally, we investigated fine‐scale patterns of genetic structure and found the spatial scale at which there was significant genetic structure varied among the regions studied. Furthermore, we discuss our results in light of other phylogeographic studies of southeastern coastal plain organisms and in relation to amphibian conservation and management.     

Turnover of labile and recalcitrant soil carbon differ in response to nitrate and ammonium deposition in an ombrotrophic peatland

The effects of 4 years of simulated nitrogen deposition, as nitrate (NO₃^?) and ammonium (NH₄⁺), on microbial carbon turnover were studied in an ombrotrophic peatland. We investigated the mineralization of simple forms of carbon using MicroResp^? measurements (a multiple substrate induced respiration technique) and the activities of four soil enzymes involved in the decomposition of more complex forms of carbon or in nutrient acquisition: N‐acetyl‐glucosaminidase (NAG), cellobiohydrolase (CBH), acid phosphatase (AP), and phenol oxidase (PO). The potential mineralization of labile forms of carbon was significantly enhanced at the higher N additions, especially with NH₄⁺ amendments, while potential enzyme activities involved in breakdown of more complex forms of carbon or nutrient acquisition decreased slightly (NAG and CBH) or remained unchanged (AP and PO) with N amendments. This study also showed the importance of distinguishing between NO₃^? and NH₄⁺ amendments, as their impact often differed. It is possible that the limited response on potential extracellular enzyme activity is due to other factors, such as limited exposure to the added N in the deeper soil or continued suboptimal functioning of the enzymes due to the low pH, possibly via the inhibitory effect of low phenol oxidase activity.     

New microsatellite loci for the European bushcricket,Ephippiger ephippiger (Orthoptera: Tettigoniidae)

Ephippiger ephippiger is a model organism for studies of sexual selection and phylogeography but little is known about fine‐scale population structure. Available microsatellite loci have null allele problems so we used an enrichment technique to isolate 21 new microsatellite loci for E. ephippiger. We present primer pairs for 10 polymorphic loci (3–11 alleles per locus). Observed heterozygosities at polymorphic loci ranged from 0.118 to 0.787, but several were significantly lower than expected.     

Effects of experimental drought on soil respiration and radiocarbon efflux from a temperate forest soil

Soil moisture affects microbial decay of SOM and rhizosphere respiration (RR) in temperate forest soils, but isolating the response of soil respiration (SR) to summer drought and subsequent wetting is difficult because moisture changes are often confounded with temperature variation. We distinguished between temperature and moisture effects by simulation of prolonged soil droughts in a mixed deciduous forest at the Harvard Forest, Massachusetts. Roofs constructed over triplicate 5 × 5 m² plots excluded throughfall water during the summers of 2001 (168 mm) and 2002 (344 mm), while adjacent control plots received ambient throughfall and the same natural temperature regime. In 2003, throughfall was not excluded to assess the response of SR under natural weather conditions after two prolonged summer droughts. Throughfall exclusion significantly decreased mean SR rate by 53 mg C m^?2 h^?1 over 84 days in 2001, and by 68 mg C m^?2 h^?1 over 126 days in 2002, representing 10–30% of annual SR in this forest and 35–75% of annual net ecosystem exchange (NEE) of C. The differences in SR were best explained by differences in gravimetric water content in the Oi horizon (r²=0.69) and the Oe/Oa horizon (r²=0.60). Volumetric water content of the A horizon was not significantly affected by throughfall exclusion. The radiocarbon signature of soil CO₂ efflux and of CO₂ respired during incubations of O horizon, A horizon and living roots allowed partitioning of SR into contributions from young C substrate (including RR) and from decomposition of older SOM. RR (root respiration and microbial respiration of young substrates in the rhizosphere) made up 43–71% of the total C respired in the control plots and 41–80% in the exclusion plots, and tended to increase with drought. An exception to this trend was an interesting increase in CO₂ efflux of radiocarbon‐rich substrates during a period of abundant growth of mushrooms. Our results suggest that prolonged summer droughts decrease primarily heterotrophic respiration in the O horizon, which could cause increases in the storage of soil organic carbon in this forest. However, the C stored during two summers of simulated drought was only partly released as increased respiration during the following summer of natural throughfall. We do not know if this soil C sink during drought is transient or long lasting. In any case, differential decomposition of the O horizon caused by interannual variation of precipitation probably contributes significantly to observed interannual variation of NEE in temperate forests.     

A comparative study of cannibalism and predation in seven species of flour beetle

Abstract 1. The present study quantified egg and pupal cannibalism, and interspecific predation on eggs and pupae, by larvae and adults of seven species of flour beetle (Tribolium spp.) under laboratory conditions: T. anaphe, T. brevicornis, T. castaneum, T. confusum, T. destructor, T. freemani, and T. madens. 2. Variation among species in cannibalism and predation propensities did not reflect taxonomic affinities within the genus, indicating that these behaviours were shaped by ecology at species level. 3. Within species, larvae and adults displayed different propensities for cannibalism and predation, leading to the conclusion that these behaviours evolve independently in the two life stages. 4. All species behaved as intraguild predators to some degree, especially in the adult stage. 5. Three general patterns of cannibalism and predation were described by principal component mapping and cluster analysis. 6. The first group comprised three cosmopolitan pest species that were more voracious as adults than as larvae: T. castaneum, T. confusum, and T. destructor. It is proposed that stored product environments select for high adult voracity because the costs associated with emigration from such rare, but resource‐rich, habitats intensifies interference competition among adults. 7. The second group consisted of species that inhabit natural environments and that were more voracious as larvae: T. anaphe, T. freemani, and T. madens. Habitats for these species are probably numerous, but generally poor in quality, a situation that intensifies larval competition, while favouring earlier adult emigration. 8. The largest species, T. brevicornis, demonstrated inconsistent voracity between life stages and was the only species with chemically defended pupae. 9. It is proposed that consumption of eggs provides primarily nutritional benefits, whereas consumption of pupae has a more important role in interference competition.