Egg mass and incubation period allometry in birds and reptiles: effects of phylogeny

Here we test the hypothesis that the relationship between egg mass at oviposition (IEM) and incubation period ( I _p) is a function of the taxonomic relatedness of bird and reptile species. Allometric relationships between IEM and I _p were examined for 1525 bird species and 201 reptilian species. Treating species as independent data revealed the allometric exponent linking I _p to IEM to be 0.234 for birds and 0.138 for reptiles. However, ANCOVA revealed that within both birds and reptiles the elevation and slope of the regression lines were dependent on the taxonomic order studied, indicating that the exponents were confounded by the phylogenetic relatedness of species. Thus, allometric exponents were recalculated based on the method of comparative analysis using independent contrasts. This technique revealed that the allometric exponent in both birds and reptiles was confounded by phylogeny. In birds the allometric relationship between I _p and IEM was almost halved to 0.122, whereas in reptiles the exponent increased to 0.185. Importantly, the results demonstrate that some results of allometric analyses can be artefacts of the method of analysis of the dataset. That for bird eggs I _p is not determined in large part by egg mass allows new questions to be posed regarding the ecological and physiological factors affecting the length of incubation, and hence rates of embryonic growth, for different taxa and habitats.     

An exploration of differences in the scaling of life history traits with body mass within reptiles and between amniotes

Allometric relationships linking species characteristics to body size or mass (scaling) are important in biology. However, studies on the scaling of life history traits in the reptiles (the nonavian Reptilia) are rather scarce, especially for the clades Crocodilia, Testudines, and Rhynchocephalia (single extant species, the tuatara). Previous studies on the scaling of reptilian life history traits indicated that they differ from those seen in the other amniotes (mammals and birds), but so far most comparative studies used small species samples and also not phylogenetically informed analyses. Here, we analyzed the scaling of nine life history traits with adult body mass for crocodiles (n = 22), squamates (n = 294), turtles (n = 52), and reptiles (n = 369). We used for the first time a phylogenetically informed approach for crocodiles, turtles, and the whole group of reptiles. We explored differences in scaling relationships between the reptilian clades Crocodilia, Squamata, and Testudines as well as differences between reptiles, mammals, and birds. Finally, we applied our scaling relationships, in order to gain new insights into the degree of the exceptionality of the tuatara's life history within reptiles. We observed for none of the life history traits studied any difference in their scaling with body mass between squamates, crocodiles, and turtles, except for clutch size and egg weight showing small differences between these groups. Compared to birds and mammals, scaling relationships of reptiles were similar for time‐related traits, but they differed for reproductive traits. The tuatara's life history is more similar to that of a similar‐sized turtle or crocodile than to a squamate.     

THE PHYSIOLOGICAL ECOLOGY OF REPTILIAN EGGS AND EMBRYOS. AND THE EVOLUTION OF VIVIPARITY WITHIN THE CLASS REPTILIA

1. Eggs of Crocodilia and Chelonia, like those of birds, have a pair of egg membranes separating a thick layer of albumen from the calcareous shell. In contrast, eggs of oviparous Lepidosauria have only a single shell membrane, upon which relatively small amounts of calcium carbonate are deposited; and the volume of albumen in eggs is extraordinarily small at the time of oviposition. 2. With the possible exception of certain geckos and some chelonians, eggs of oviparous reptiles seem always to absorb water from the substrate during the course of normal incubation. In so far as the rate of water absorption exceeds the rate of water loss by transpiration from exposed surfaces, the eggs swell during incubation. The term ‘cleidoic’ cannot be used to describe eggs of this type. 3. Embryos of lizards and snakes influence the water potential of extra-embryonic fluids contained within their eggs, thereby maintaining or increasing the gradient in water potential that drives water absorption. 4. Embryos of Crocodilia and Chelonia obtain a substantial portion of the calcium used in ossification of skeletal elements from the inner surfaces of the eggshell. In contrast, embryonic lizards and snakes draw upon extensive reserves of calcium present in the yolk, and obtain little (if any) calcium from the eggshell. 5. All reptilian embryos seem to produce substantial quantities of urea as a detoxification product of protein catabolism. Contrary to expectation, uricotelism may not be common among reptilian embryos, even in those few instances where development takes place within a hard, calcareous egg. 6. In eggs of Crocodilia and Chelonia, respiratory gases seem to pass by diffusion through pores in the calcareous eggshell and through spaces between the fibres of the pair of egg membranes. No pores have been observed in the shell of lepidosaurian eggs, and so gases presumably diffuse between the fibres of the single (multilayered) shell membrane. 7. Metabolism of reptilian embryos is temperature-dependent, as is true for most ectothermic organisms. For each species, there appears to be a particular temperature at which embryonic development proceeds optimally, and departures from this optimum elicit increases in developmental anomalies and/or embryonic mortality. 8. Viviparity has evolved on numerous occasions among species of the Squamata, but seemingly never among Crocodilia or Chelonia. Since the evolution of viviparity entails a progressive reduction in the eggshell, only those organisms whose embryos do not depend upon the eggshell as a source of calcium may have the evolutionary potential to become viviparous. 9. Evolutionary transitions from oviparity to viviparity could have been driven by selection related to (i) thermal benefits to embryos consequent upon retention of eggs within the body of a parent capable of behavioural thermoregulation; (ii) protection of the eggs from nest predators and/or soil microbes; and (iii) more effective exploitation of a seasonal food resource by early emerging young.     

应用混合模型和分位数回归分析东黄海小黄鱼肥满度空间异质性

基于2008-2010年黄海南部近海(SYS)、东海北部外海(NECS)和东海中部近海(MECS)小黄鱼体长和体质量数据,采用均值回归和分位数回归模型,解析了小黄鱼幼鱼和成鱼群体体长-体质量关系的空间变异.结果表明: 协方差模型和线性混合模型的残差标准误基本一致,线性模型残差标准误最高.从线性混合模型对特定区域和总体区域平均体质量计算的相对比值来看,SYS和NECS幼鱼群体的平均体质量高于总体平均值,但MECS低于总体平均值;成鱼群体则为NECS平均体质量高于总体平均值,MECS和SYS低于总体平均值.分位回归估计的肥满度和异速生长指数结果显示,幼鱼群体在不同分位的估计参数呈显著变化,SYS异速生长指数均值为2.85,在0.1~0.95分位的估计值变化范围为2.63~2.96.MECS和NECS参数估计值和置信区间在各分位数呈异质性变化,低分位时,NECS估计值在3个调查区域中最低,MECS最高;高分位时,MECS和NECS均高于SYS.对低分位0.25、中分位0.5和高分位0.75分位数的异速体长体质量关系的方差分析结果显示,低分位和高分位数之间体长 体质量关系极为显著(0.25∶0.75,F=6.38,df=1737,P<0.01),低分位数和中分位数之间为显著(0.25∶0.5,F=2.35,df=1737,P=0.039),中分位数和高分位数之间接近显著(0.5∶0.75,F=2.21,df=1737,P=0.051).成鱼群体SYS异速生长指数均值为3.01,在0.1~0.95分位的估计值变化范围为2.77~3.10.低分位和高分位数之间体长 体质量关系差异达到显著水平(0.25∶0.75,F=3.31,df=2793,P=0.01),低分位和中分位之间差异不显著(0.25∶0.5,F=0.98,df=2793,P=0.43),而高分位和中分位之间则差异极显著(0.5∶0.75,F=3.56,df=2793,P<0.01).     

Genomic V exons from whole genome shotgun data in reptiles

Reptiles and mammals diverged over 300 million years ago, creating two parallel evolutionary lineages amongst terrestrial vertebrates. In reptiles, two main evolutionary lines emerged: one gave rise to Squamata, while the other gave rise to Testudines, Crocodylia, and Aves. In this study, we determined the genomic variable (V) exons from whole genome shotgun sequencing (WGS) data in reptiles corresponding to the three main immunoglobulin (IG) loci and the four main T cell receptor (TR) loci. We show that Squamata lack the TRG and TRD genes, and snakes lack the IGKV genes. In representative species of Testudines and Crocodylia, the seven major IG and TR loci are maintained. As in mammals, genes of the IG loci can be grouped into well-defined IMGT clans through a multi-species phylogenetic analysis. We show that the reptilian IGHV and IGLV genes are distributed amongst the established mammalian clans, while their IGKV genes are found within a single clan, nearly exclusive from the mammalian sequences. The reptilian and mammalian TRAV genes cluster into six common evolutionary clades (since IMGT clans have not been defined for TR). In contrast, the reptilian TRBV genes cluster into three clades, which have few mammalian members. In this locus, the V exon sequences from mammals appear to have undergone different evolutionary diversification processes that occurred outside these shared reptilian clans. These sequences can be obtained in a freely available public repository (http://vgenerepertoire.org).     

The relationship between body mass and relative investment in testes mass in amniotes and other vertebrates

In terrestrial placental mammals, there is a well‐known negative allometric relationship between body mass and relative investment in testes mass. Such a negative relationship means that males of relatively monogamous small species invest proportionately more in their reproductive tissues than males of more polyandrous larger species. The selective pressure responsible for this relationship remains unclear and is it not known if this is a general allometric relationship that is similar across all vertebrate lineages. To investigate this, we conducted the first comparison of relationships between body mass and testes mass (using percentage testes mass as the dependent variable) across a variety of vertebrate groups. In all amniote lineages examined, the allometric relationship between body mass and testes mass was relatively strong and negative. We show, for the first time, that reptiles, birds and terrestrial placental mammals followed the same allometric relationship and, contrary to previous expectations, this relationship is sigmoidal rather than linear. Within this data set, there was no significant difference between this general amniote relationship and any of the 13 orders of reptiles, birds and terrestrial placental mammals examined. As a result, we propose that a sigmoidal relationship should be considered the default assumption for the form of the body mass – testes mass relationship within the amniote lineage. However, we also identify significant differences within some additional mammal groups (marsupials, bats and cetaceans). In each of these cases, only some sub‐groupings differed significantly from the general amniote relationship. In contrast to the amniotes, the relationship is relatively weak and positive in teleost fish and frogs suggesting that a negative allometric relationship is not universal in vertebrates. We explore whether variation in the body mass – testes mass relationships can be linked to sperm competition or a variety of ecological characteristics, either for amniotes in particular or vertebrates in general.     

Scaling of maximal oxygen uptake by lower leg muscle volume in boys and men.

The aim of this study was to critically examine the influence of body size on maximal oxygen uptake (VO2 max) in boys and men using body mass (BM), estimated fat-free mass (FFM), and estimated lower leg muscle volume (Vol) as the separate scaling variables. VO2 max and an in vivo measurement of Vol were assessed in 15 boys and 14 men. The FFM was estimated after percentage body fat had been predicted from population-specific skinfold measurements. By using nonlinear allometric modeling, common body size exponents for BM, FFM, and Vol were calculated. The point estimates for the size exponent (95% confidence interval) from the separate allometric models were: BM 0.79 (0.53-1.06), FFM 1.00 (0.78-1.22), and Vol 0.64 (0.40-0.88). For the boys, substantial residual size correlations were observed for VO2 max/BM0.79 and VO2 max/FFM1.00, indicating that these variables did not correctly partition out the influence of body size. In contrast, scaling by Vol0.64 led to no residual size correlation in boys or men. Scaling by BM is confounded by heterogeneity of body composition and potentially substantial differences in the mass exponent between boys and men. The FFM is precluded as an index of involved musculature because Vol did not represent a constant proportion of FFM [Vol proportional, variantFFM1.45 (95% confidence interval, 1.13-1.77)] in the boys (unlike the men). We conclude that Vol, as an indicator of the involved muscle mass, is the most valid allometric denominator for the scaling of VO2 max in a sample of boys and men heterogeneous for body size and composition.     

Intraspecific allometry of standard metabolic rate in green iguanas, Iguana iguana

To study the allometric relationship between standard metabolic rate and body mass (mass range 16-3627 g) in green iguanas, Iguana iguana (n=32), we measured rates of oxygen consumption (V(O(2))) at 30 degrees C during scotophase. The relationship could be described as: V(O(2))(ml h(-1))=0.478W(0.734). The resulting mass exponent was similar to the 3/4 power commonly used in interspecific curves (P>0.05), but differed from a proposed intraspecific value of 2/3 (P<0.05). The mass exponents of male (n=8) and female (n=11) iguanas did not differ (P>0.05). The mass adjusted V(O(2)) was higher than predicted from generalized squamate curves. The mean mass exponent of intra-individual allometric equations of iguanas (n=7) at varying masses during ontogeny did not differ from that of the pooled equation, indicating that scaling of V(O(2)) is similar for both between and within individuals. Thermal acclimation, compensatory changes in V(O(2)) with prolonged exposure to a constant temperature, was not observed in juvenile iguanas (n=11) between 1 and 5 weeks of acclimation at 30 degrees C.     

Basal metabolic rates in mammals: taxonomic differences in the allometry of BMR and body mass

No single equation adequately describes the allometric relation between body mass and BMR for mammals. Least squares regression of log-transformed data for 248 eutherian species results in a line with a slope (-0.30) significantly different from that of Kleiber's line (-0.25). Interordinal comparisons of least squares regressions of log-transformed BMR and mass suggest that the Insectivora have a significantly steeper slope to their allometric relationship than do most other orders, while the non-insectivore orders are statistically homogeneous with respect to slope. With respect to elevation, Edentata have the lowest BMRs; Marsupialia, Primates and Chiroptera are indistinguishable from each other but above the edentates; Primates, Chiroptera, Rodentia, Lagomorpha and Carnivora form the next highest homogeneous grouping; and Artiodactyla have the highest BMRs, significantly greater than all but Lagomorpha and Carnivora. Analysis of intraordinal variation within the Rodentia suggests significant heterogeneity among families in BMR-mass allometry.     

Beyond the '3/4-power law': variation in the intra- and interspecific scaling of metabolic rate in animals

In this review I show that the '3/4-power scaling law' of metabolic rate is not universal, either within or among animal species. Significant variation in the scaling of metabolic rate with body mass is described mainly for animals, but also for unicells and plants. Much of this variation, which can be related to taxonomic, physiological, and/or environmental differences, is not adequately explained by existing theoretical models, which are also reviewed. As a result, synthetic explanatory schemes based on multiple boundary constraints and on the scaling of multiple energy-using processes are advocated. It is also stressed that a complete understanding of metabolic scaling will require the identification of both proximate (functional) and ultimate (evolutionary) causes. Four major types of intraspecific metabolic scaling with body mass are recognized [based on the power function R=aMb, where R is respiration (metabolic) rate, a is a constant, M is body mass, and b is the scaling exponent]: Type I: linear, negatively allometric (b<1); Type II: linear, isometric (b=1); Type III: nonlinear, ontogenetic shift from isometric (b=1), or nearly isometric, to negatively allometric (b<1); and Type IV: nonlinear, ontogenetic shift from positively allometric (b>1) to one or two later phases of negative allometry (b<1). Ontogenetic changes in the metabolic intensity of four component processes (i.e. growth, reproduction, locomotion, and heat production) appear to be important in these different patterns of metabolic scaling. These changes may, in turn, be shaped by age (size)-specific patterns of mortality. In addition, major differences in interspecific metabolic scaling are described, especially with respect to mode of temperature regulation, body-size range, and activity level. A 'metabolic-level boundaries hypothesis' focusing on two major constraints (surface-area limits on resource/waste exchange processes and mass/volume limits on power production) can explain much, but not all of this variation. My analysis indicates that further empirical and theoretical work is needed to understand fully the physiological and ecological bases for the considerable variation in metabolic scaling that is observed both within and among species. Recommended approaches for doing this are discussed. I conclude that the scaling of metabolism is not the simple result of a physical law, but rather appears to be the more complex result of diverse adaptations evolved in the context of both physico-chemical and ecological constraints.     

Foraging time and dietary intake by breeding Ross's and Lesser Snow Geese

We compared foraging times of female Ross's (Chen rossii) and Lesser Snow Geese (Chen caerulescens caerulescens) breeding at Karrak Lake, NT, Canada and examined variation due to time of day and reproductive stage. We subsequently collected female geese that had foraged for known duration and we estimated mass of foods consumed during foraging bouts. Female Ross's Geese spent more time foraging (mean % - SE =28.4ǃ.3%; P=0.0002), on average, than did female Lesser Snow Geese (21.5 - 1.4%). Foraging time by female geese differed among reproductive stages, but differences were not consistent among time periods (stage-by-time block interaction, P=0.0003). Females spent considerably more time foraging during prelaying and laying than during incubation. Ross's Geese also spent a greater percent of time feeding (83.0DŽ.8%) during incubation recesses than did Lesser Snow Geese (60.9Dž.6%). Consumption of organic matter during foraging bouts was minimal; estimated consumption averaged 9.6dž.0 and 12.4dž.6 g (mean - SE) dry mass/day before incubation and 5.9DŽ.0 and 5.7DŽ.1 g dry mass/day during incubation for Lesser Snow and Ross's Geese, respectively. Diets consisted primarily of mosses (bryophytes), Chickweed (Stellaria spp.) and Sedges (Carex spp.). Before incubation, eggshell consumption was estimated as 4.3Dž.2 and 0.4ǂ.3 g dry mass/day for Lesser Snow and Ross's Geese, respectively; neither species consumed eggshell during incubation. We conclude that eggshell from nests of previous years is likely an important source of dietary calcium used to meet mineral demands of eggshell formation at Karrak Lake. Our findings of wide disparities between foraging time and food intake indicate that results from studies that do not directly measure intake rates remain equivocal. Finally, we propose four hypotheses accounting for foraging effort that evidently yields little nutritional or energetic benefit to geese nesting at Karrak Lake.     

Molecular systematics of primary reptilian lineages and the tuatara mitochondrial genome

We provide phylogenetic analyses for primary Reptilia lineages including, for the first time, Sphenodon punctatus (tuatara) using data from whole mitochondrial genomes. Our analyses firmly support a sister relationship between Sphenodon and Squamata, which includes lizards and snakes. Using Sphenodon as an outgroup for select squamates, we found evidence indicating a sister relationship, among our study taxa, between Serpentes (represented by Dinodon) and Varanidae. Our analyses support monophyly of Archosauria, and a sister relationship between turtles and archosaurs. This latter relationship is congruent with a growing set of morphological and molecular analyses placing turtles within crown Diapsida and recognizing them as secondarily anapsid (lacking a skull fenestration). Inclusion of Sphenodon, as the only surviving member of Sphenodontia (with fossils from the mid-Triassic), helps to fill a sampling gap within previous analyses of reptilian phylogeny. We also report a unique configuration for the mitochondrial genome of Sphenodon, including two tRNA(Lys) copies and an absence of ND5, tRNA(His), and tRNA(Thr) genes.     

Analysis of heart rate in developing bird embryos: effects of developmental mode and mass

Bird embryos may be regarded as developing in their thermo-neutral zone, at rest, and stay in the egg for a fixed period of time until hatching. It is therefore interesting to investigate if they follow the same 'rule' set for adult homeotherms, which states that, within a taxonomically or functionally defined category such as mammals or birds, the number of heart beats throughout the life span (sL) is more or less constant. This rule stems from the allometric relationships between heart rate (fH) and body mass (mB) and between sL and mB. As a step towards understanding the general allometric nature of avian embryonic physiology we analyzed the fH values of avian embryos in relation to their incubation span (sI). Data from 30 species were selected from the scientific literature for the analyses. Values obtained from invasive methods which were judged to grossly alter natural incubation conditions, or from undefined or unmatched temperature conditions were not used. These include most values obtained below the first 30% of the incubation. Also, data obtained after internal pipping were discarded since hatching activity influences them. Values for sI and egg mass (mE) as representatives of embryonic mass were also collected. Embryonic fH was normalized to 70.1-80% sI. At 20.1-30% sI it was only 85% of the value at 70.1-80% sI and increased to a plateau at about 50.1-60% sI. It was almost constant among species between 50.1 and 60% sI and pre-internal pipping (PIP) time and thus, the mean fH value between 50.1 and 60% sI and between 90.1 and 100% excluding pipped eggs (fH) was taken as a representative value for each given species. The fH (min-1) and the corresponding sI (days) values for the 30 species, scaled with mE (g) as follows: fH = 371.1.mE-0.112 and: sI = 12.29.mE+0.209. Both powers were significantly different from 0. The product of fH and sI (fH.sI), representing the total number of heartbeats throughout the incubation, scaled with mE for the entire data set as follows: fH.sI = 6.565 x 10(+6).mE+0.096, where the +0.096 power is significantly different from 0. Values for fH.sI from embryos of altricial birds tended to concentrate at the low mE end of the plot while those of the precocial ones tended towards the high end. Separate analyses showed that the mE power for the combined altricial and semi-altricial species (ASA), and the combined precocial and semi precocial species (PSP), of log fH.sI against log mE regressions, were both insignificantly different from 0. Thus, means of fH.sI for ASA and PSP were calculated. The mean ASA value of 7.27 x 10(+6) heartbeats for fH.sI, was significantly different from the mean PSP value of 10.93 x 10(+6). The difference of 3.66 x 10(-6) (33.5%) heartbeats can be attributed to either the more advanced stage of the PSP hatchlings at hatch, to the larger mE values of these hatchlings, to the difference in water fraction of the hatchlings or all. The result of a linear regression of fH.sI against the rate of sI completion (the inverse of incubation span, fI; day-1) was: fH.10(-6) = 0.205 + 3.940.sI-1. Thus, the faster is the average rate of development accomplished per day (shorter incubation) the higher is daily heart rate. Data tended to cluster such that large eggs, mostly of the PSP type with relatively low fH, complete 2-4% of their incubation per day, while small, ASA type eggs with relatively high fH, complete 6-8% of their incubation time per day. We conclude that, at this stage of knowledge, the data is insufficient to resolve whether the different modes of hatch stage alone can explain differences in the total number of heartbeats throughout embryonic life among all bird species, or egg mass and water content differences contribute variability. This should be investigated on a larger sample of species in more depth.     

The relationship of egg size and incubation temperature to embryonic development time in univoltine and multivoltine aquatic insects

1. We used published data to investigate the combined influence of egg size and incubation temperature on embryonic development time for a broad assortment of aquatic insects at four different incubation temperatures (10, 15, 20 and 25 °C).
2. Embryonic development time (EDT) was positively correlated with egg size at each of the four temperatures, but with different relationships for univoltine and multivoltine aquatic insects. The relationships of embryonic development time to egg size expressed in degree-days did not significantly differ in slope ( P >0.50) or intercept ( P >0.05) for either univoltine or multivoltine aquatic insects at each of the four temperatures.
3. The relationship of embryonic development time (degree-days) to egg mass in multivoltine aquatic insects (EDT=885×0.19, P <0.0001, r 2=0.48) is similar in slope and intercept to that for other oviparous animals (i.e., zooplankton, fish, amphibians and reptiles), and to the relationship of embryonic development time to neonate mass in mammals. Univoltine species on average require 3–5 times longer to develop (EDT=14190×0.29, P <0.001, r 2=0.29) than most other animals of equivalent egg mass, but the relationship of embryonic development time to egg mass is similar in slope to that of most other animals. Together, these relationships provide a basis for evaluating differences in embryonic development time among aquatic insects.     

Ecogeographical and phylogenetic effects on craniofacial variation in macaques

The widespread and complex ecogeographical diversity of macaques may have caused adaptive morphological convergence among four phylogenetic subgroups, making their phylogenetic relationships unclear. We used geometric morphometrics and multivariate analyses to test the null hypothesis that craniofacial morphology does not vary with ecogeographical and phylogenetic factors. As predicted by Bergmann's rule, size was larger for the fascicularis and sinica groups in colder environments. No clear size cline was observed in the silenus and sylvanus groups. An allometric pattern was observed across macaques, indicating that as size increases, rounded faces become more elongated. However, the elevation was differentiated within each of the former two groups and between the silenus and sylvanus groups, and the slope decreased in each of the two northern species of the fascicularis group. All allometric changes resulted in the similar situation of the face being more rounded in animals inhabiting colder zones and/or in animals having a larger body size than that predicted from the overarching allometric pattern. For non‐allometric components, variations in prognathism were significantly correlated with dietary differences; variations in localized shape components in zygomatics and muzzles were significantly correlated with phylogenetic differences among the subgroups. The common allometric pattern was probably influenced directly or indirectly by climate‐related factors, which are pressures favoring a more rounded face in colder environments and/or a more elongated face in warmer environments. Allometric dissociation could have occurred several times in Macaca even within a subgroup because of their wide latitudinal distributions, critically impairing the taxonomic utility of craniofacial elongation. Am J Phys Anthropol 154:27–41, 2014. © 2014 Wiley Periodicals, Inc.     

A developmental synapomorphy of squamate reptiles

The reptilian clade Squamata is defined primarily by osteological synapomorphies, few of which are entirely unambiguous. Studies of developing squamate eggs have revealed a uniquely specialized feature not known to occur in any other amniotes. This feature—the yolk cleft/isolated yolk mass complex—lines the ventral hemisphere of the egg. During its formation, extraembryonic mesoderm penetrates the yolk and an exocoelom (the yolk cleft [YC]) forms in association with it, cutting off a thin segment of yolk (the “isolated yolk mass” [IYM]) from the main body of the yolk. The YC–IYM complex has been observed and described in more than 65 squamate species in 12 families. In viviparous species, it contributes to the “omphaloplacenta,” a type of yolk sac placenta unique to squamates. The only squamates known to lack the IYM are a few highly placentotrophic skinks with minuscule eggs, viviparous species in which it clearly has been lost. Given its absence in mammals, chelonians, crocodylians, and birds, the YC–IYM complex warrants recognition as a developmental synapomorphy of the squamate clade. As in extant viviparous lizards and snakes, the YC–IYM complex presumably contributed to the placenta of extinct viviparous squamates.     

An identification of invariants in life history traits of amphibians and reptiles

While many morphological, physiological, and ecological characteristics of organisms scale with body size, some do not change under size transformation. They are called invariant. A recent study recommended five criteria for identifying invariant traits. These are based on that a trait exhibits a unimodal central tendency and varies over a limited range with body mass (type I), or that it does not vary systematically with body mass (type II). We methodologically improved these criteria and then applied them to life history traits of amphibians, Anura, Caudata (eleven traits), and reptiles (eight traits). The numbers of invariant traits identified by criteria differed across amphibian orders and between amphibians and reptiles. Reproductive output (maximum number of reproductive events per year), incubation time, length of larval period, and metamorphosis size were type I and II invariant across amphibians. In both amphibian orders, reproductive output and metamorphosis size were type I and II invariant. In Anura, incubation time and length of larval period and in Caudata, incubation time were further type II invariant. In reptiles, however, only number of clutches per year was invariant (type II). All these differences could reflect that in reptiles body size and in amphibians, Anura, and Caudata metamorphosis (neotenic species go not through it) and the trend toward independence of egg and larval development from water additionally constrained life history evolution. We further demonstrate that all invariance criteria worked for amphibian and reptilian life history traits, although we corroborated some known and identified new limitations to their application.     

上海辰山植物园不同生活型木本植物枝叶大小关系的比较

植物枝叶生长普遍存在显著的正相关关系，决定了植物构型塑造和生物量分配。以上海辰山植物园149种木本植物为对象，通过测定顶枝上70 cm长枝条的直径、叶面积及其生物量，比较相同生境条件下不同生活型木本植物的枝叶大小关系。结果表明，枝条截面积与总叶面积间呈异速生长关系（a=1.148 6，CI=1.000 6~1.302 3），枝条干重与叶干重间呈等速生长关系（a=1.054 2，CI=0.921 3~1.205 6），不同生活型均具有相同的斜率系数a。不同生活型的异速生长常量b（y轴截距）存在显著差异，相同枝条截面积下落叶乔木比常绿乔木和常绿灌木具有更大的叶面积，相同枝条干重下常绿乔木和落叶乔木比常绿灌木具有更大的叶干重。这可能与不同生活型木本植物水分竞争效率和叶建成成本差异有关。