Anamorphosis in millipedes (Diplopoda)—the present state of knowledge with some developmental and phylogenetic considerations

A review of the postembryonic development of millipedes (Diplopoda) is given, based mainly on published information. Original observations are, however, also included. Millipedes hatching from the pupoid usually have three pairs of legs; during their postembryonic growth they acquire more segments and more legs. This process is known as anamorphosis. Three types of anamorphosis are recognized. In euanamorphosis, every moult is accompanied by addition of new segments, even after the attainment of sexual maturity. In hemianamorphosis, the addition of new segments goes on until a certain stadium, and further moults take place without addition of segments. In teloanamorphosis, the addition of segments stops at a certain stadium (the adult, and ultimate, stadium) after which no further moults occur. Available information on anamorphosis in each of the millipede orders is reviewed. General patterns are emphasized, but variations are also considered in detail. It is shown that the so-called ‘law of anamorphosis’ is valid only for the ‘ring-forming’ millipedes (Merocheta and Juliformia) in which tergites, pleurites, and sternites of each diplosegment are firmly fused into a complete ‘ring’, and for some other forms (Polyzoniida, Chordeumatida), where there is a constant relationship between rings and legs. The chapter on the order Julida is particularly detailed and includes discussions of patterns in the variation and a section on periodomorphosis. The general chapter on developmental patterns includes inter alia an interpretation of the variations in millipede anamorphosis in terms of the ‘biometabolic modi’ of Remane. The hypothetical ancestral millipede is shown to have developed by hemianamorphosis. Euanamorphosis was acquired by the ancestral species of Helminthomorpha. Within this clade, Chordeumatida and Merocheta have secondarily become teloanamorphic, whereas some Juliformia seem to have returned to hemianamorphosis. The contrasting principles of elongation and contraction, subject of much debate among diplopodologists, are shown both to have played a role in the course of millipede evolution.     

Systematic assessment of terrestrial biogeochemistry in coupled climate–carbon models

With representation of the global carbon cycle becoming increasingly complex in climate models, it is important to develop ways to quantitatively evaluate model performance against in situ and remote sensing observations. Here we present a systematic framework, the Carbon‐LAnd Model Intercomparison Project (C‐LAMP), for assessing terrestrial biogeochemistry models coupled to climate models using observations that span a wide range of temporal and spatial scales. As an example of the value of such comparisons, we used this framework to evaluate two biogeochemistry models that are integrated within the Community Climate System Model (CCSM) – Carnegie‐Ames‐Stanford Approach′ (CASA′) and carbon–nitrogen (CN). Both models underestimated the magnitude of net carbon uptake during the growing season in temperate and boreal forest ecosystems, based on comparison with atmospheric CO₂ measurements and eddy covariance measurements of net ecosystem exchange. Comparison with MODerate Resolution Imaging Spectroradiometer (MODIS) measurements show that this low bias in model fluxes was caused, at least in part, by 1–3 month delays in the timing of maximum leaf area. In the tropics, the models overestimated carbon storage in woody biomass based on comparison with datasets from the Amazon. Reducing this model bias will probably weaken the sensitivity of terrestrial carbon fluxes to both atmospheric CO₂ and climate. Global carbon sinks during the 1990s differed by a factor of two (2.4 Pg C yr^?1 for CASA′ vs. 1.2 Pg C yr^?1 for CN), with fluxes from both models compatible with the atmospheric budget given uncertainties in other terms. The models captured some of the timing of interannual global terrestrial carbon exchange during 1988–2004 based on comparison with atmospheric inversion results from TRANSCOM (r=0.66 for CASA′ and r=0.73 for CN). Adding (CASA′) or improving (CN) the representation of deforestation fires may further increase agreement with the atmospheric record. Information from C‐LAMP has enhanced model performance within CCSM and serves as a benchmark for future development. We propose that an open source, community‐wide platform for model‐data intercomparison is needed to speed model development and to strengthen ties between modeling and measurement communities. Important next steps include the design and analysis of land use change simulations (in both uncoupled and coupled modes), and the entrainment of additional ecological and earth system observations. Model results from C‐LAMP are publicly available on the Earth System Grid.