Delimitation of a continuous morphological character with unknown prior membership: application of a finite mixture model to classify scapular setae of Abacarus panticis

Unambiguous classification is a prerequisite for the study of polymorphism, but accurate delimitation of continuous morphological characters can be challenging. Finite mixture modeling is a rigorous and flexible approach for delimiting continuous variables with unknown prior membership, but its application to morphological studies remains limited. In this study, the lengths of scapular setae of the eriophyoid mite Abacarus panticis Keifer collected from 18 sites in Taiwan were used as an example to evaluate the eligibility of finite mixture models. We then tested the hypothesis that longer scapular setae can facilitate dispersal. Lastly, we investigate morphological variation in various seta morphs by geometric morphometric techniques. Finite mixture models can satisfactorily classify scapular setae of A. panticis into long and short seta morphs. Abacarus panticis of the long morph only occurred in five sites whereas the short seta morph existed in all study sites. Geometric morphometric analyses revealed a more elongated coxal area in individuals of long morph than in those of short morph. Because the short morph is more widespread in geographical distribution than the long morph, longer scapular setae seem unlikely a specialized adaptation for dispersal. Further studies should capitalize on the finite mixture model in the delimitation of continuous morphological characters.     

Body size and craniometry of the herb field mouse from Lithuania in the context of species range

We present morphometric and craniometric measurements of the herb field mouse (Apodemus uralensis) from Lithuania and analyze variation of body and skull size across species range. We suppose species is characterized by high size variability, not following Bergmann’s or Murphy’s rules. Preliminary, distinct size differences have been registered in the eastern and southern edges of the distribution range, with these populations having largest individuals according to average body and skull size. In terms of tail length and condylobasal length of the skull, Lithuanian mice on the north-western edge of the species range are among the largest, but in terms of body weight, body length, zygomatic skull width and the length of maxillary toothrow, adult A. uralensis from Lithuania are small and correspond to those from populations on the western edge of the range. The relative skull width (ratio of zygomatic skull width to condylobasal length) of Lithuanian A. uralensis is the smallest across the entire range. In A. uralensis from Lithuania, sex dimorphism is weakly expressed, with hind foot length and postorbital constriction larger in adult males, while the height of the mandibula and length of the mandibular diastema is larger in adult females. Juvenile and subadult A. uralensis from Lithuania differ in body weight, but not in size.     

Environmental Correlates of Body Mass in True Lemurs (<Emphasis Type="Italic">Eulemur</Emphasis> spp.)

An organism’s body size is intrinsically related to its metabolic requirements, life history profile, and ecological niche. Previous work in primates generally, and lemurs specifically, has shown that body size often correlates with ecological parameters related to temperature and energy availability in the environment, although other studies indicate the absence of any such patterns in lemurs. Here we test hypotheses that predict that body mass in Eulemur should covary with 1) overall food availability or resource seasonality and/or 2) temperature, i.e., Bergmann’s rule. We use data from 722 wild true lemurs to identify population-specific body mass for 27 populations representing 11 of the 12 described Eulemur species, and derive climatic data for each population from the WorldClim database. We use phylogenetic generalized least squares models to evaluate these hypotheses and find that body mass significantly negatively correlates with annual mean temperature and positively correlates with standard deviation of temperature, but does not significantly correlate with annual rainfall or number of dry months. These results indicate that body mass distribution across populations in Eulemur is consistent with Bergmann’s rule, but does not track resource seasonality as seen in other lemurs, e.g., Propithecus. We suggest that the fact that body mass in various lemur radiations correlates with different environmental variables may result in these patterns being obscured when such taxa are combined in a single analysis. This may help explain why some previous work has found little evidence of ecogeographical correlates with body mass across all Lemuriformes.     

Testing Bergmann’s rule and the Rosenzweig hypothesis with craniometric studies of the South American sea lion

We tested the validity of Bergmann’s rule and Rosenzweig’s hypothesis through an analysis of the geographical variation of the skull size of Otaria flavescens along the entire distribution range of the species (except Brazil). We quantified the sizes of 606 adult South American sea lion skulls measured in seven localities of Peru, Chile, Uruguay, Argentina, and the Falkland/Malvinas Islands. Geographical and environmental variables included latitude, longitude, and monthly minimum, maximum, and mean air and ocean temperatures. We also included information on fish landings as a proxy for productivity. Males showed a positive relationship between condylobasal length (CBL) and latitude, and between CBL and the six temperature variables. By contrast, females showed a negative relationship between CBL and the same variables. Finally, female skull size showed a significant and positive correlation with fish landings, while males did not show any relationship with this variable. The body size of males conformed to Bergmann’s rule, with larger individuals found in southern localities of South America. Females followed the converse of Bergmann’s rule at the intraspecific level, but showed a positive relationship with the proxy for productivity, thus supporting Rosenzweig’s hypothesis. Differences in the factors that drive body size in females and males may be explained by their different life-history strategies. Our analyses demonstrate that latitude and temperature are not the only factors that explain spatial variation in body size: others such as food availability are also important for explaining the ecogeographical patterns found in O. flavescens.     

Potential effects of El Niño-South Oscillation (ENSO) on growth of the American crocodile, Crocodylus acutus (Crocodylia: Crocodilydae) in captivity

Temperature affects growth rate of crocodiles. However, no information exists about the effects of temperature by El Niño-South Oscillation (ENSO) on crocodiles. In this paper, we present information about the effect of ENSO on total length and body mass of Crocodylus acutus in captivity during 1997–2001. We did not observe differences in total length among years, but we did so in body mass. Furthermore, we observed that warm episodes of ENSO were associated with a higher average total length and cold episodes represented a higher average of body mass. Sea surface temperature (SST) was significantly related with total length; however, the relationship between SST and body mass was unclear. We suggest that ENSO effects on growth rates of crocodiles need to be considered as an important factor on the management of captive populations.     

Climate,isolation and intraspecific competition affect morphological traits in an endangered steppe bird,the Dupont’s Lark Chersophilus duponti

Capsule: We assess biometric variation in the threatened Dupont’s Lark along a wide fraction of its distribution area analysing the largest data set for this bird species available to date, comprising a 28-year period and including birds captured in Spain, Morocco and Tunisia.

Aims: To analyse Dupont’s Lark morphology evaluating five potential sources of divergence: (I) sexual dimorphism, (II) macro-regional differences, (III) climate, (IV) isolation and (V) intraspecific competition.

Methods: Multivariate analysis was used to summarize biometric data. Sexual dimorphism and macro-regional divergence were assessed by generalized linear mixed models. Climate, isolation and intraspecific competition effects on phenotype were explored by means of model averaging.

Results: Sexes differed in wing shape suggesting a sexual selection pressure on males for aerial displays. Males showed longer bills after controlling for body size. We found an increasing Spain?Conclusion: We found convincing results for wing morphology variation in relation to intrasexual competition, as males seem to be subjected to a sexual selection pressure for aerial display and this adaptation strengthens when intraspecific competition increases. We also found solid support for Allen’s rule, with results suggesting that bill size plays an important role in the bird’s thermoregulation, which does not occur in the case of tarsus. Although Bergmann’s rule is not supported in relation to minimum annual temperature, we found a relationship with thermal conditions in the breeding season, as well as water availability.     

Body size varies with abundance,not climate,in an amphibian population

Body size variation among animals has many possible correlates, temporal as well as geographic. Adult male body size was analysed over the course of 23 yr (1989–2011) in a population of Fowler’s toads Anaxyrus fowleri at Long Point, Ontario. We used an information theoretic approach to identify the most likely models to explain body length variation in relation to abundance, age and environmental variables, including temperature. Male toads overall averaged 53.6 ± 0.1 (SE) mm (n = 1976) but average body length from year to year varied from 50.9 ± 0.2 to 61.4 ± 1.3 mm (n = 23 yr), a difference of 18.7%. Abundance was the only variable significantly correlated with body size variation (R²= 0.713, p = < 0.001). A significant 10‐yr trend in increased body size (R²= 0.874, p = < 0.001) was coincident with a previously detected negative trend in abundance. A 0.05°C yr^?1 increase in environmental temperature over the course of our study was not significantly correlated with the toads’ body size. Body size variation in these toads is likely related to density‐dependent resource availability for growth in the terrestrial stage. Temporal changes in average body size within populations in relation to density may be a significant component of phenotypic variation.     

An Intercontinental Analysis of Climate-Driven Body Size Clines in Reptiles: No Support for Patterns, No Signals of Processes

Climatic gradients impose clinal selection on animal ecological and physiological performance, often promoting geographic body size clines. Bergmann’s rule predicts that body size increases with decreasing environmental temperatures given the need to retain body-heat through adjustments of body-mass-to-surface-area ratio. This prediction generally holds for endotherms, but remains controversial for ectotherms. An alternative interpretation, the ‘resource rule’, suggests that food abundance, primary productivity and precipitation (which, unlike temperature, do not necessarily correlate with geography), drive body size clines. We investigate geographic variation in body size within 65 species of lizards and snakes (squamates) based on an intercontinental dataset (6,500+ specimens belonging to 56 Israeli species, and multiple populations of nine Liolaemus species from Argentina and Chile). Bergmann’s rule is only rarely supported by our data (in four species, 6 %), whereas six species (9 %) follow its converse (hence, it is unsupported in 94 % of cases). Similarly, size increases with resource abundance in only 12 species (18 %). Therefore, although neither of the rules is supported, factors suggested by the resource rule are better predictors of body size than temperature. Surprisingly, we show that some measures of the extent of a species’ climatic envelope do not affect the likelihood of it showing a size-climate relationship. We conclude that negative size-temperature associations are an exception rather than a generality among squamates.     

Development time and size-related traits in the oriental blowfly, Chrysomya megacephala along a latitudinal gradient from China

Organisms can respond to variation in temperature through the direct effect of temperature on phenotypes (phenotypic plasticity), or through long-term adaptation to temperature (and thus evolution of either mean size or thermal reaction norm). We examined the effects of various temperatures (of 20 and 30 °C) on development time, adult body size (body length and body width) and pre-adult survivorship in six populations of Chrysomya megacephala, collected at different latitudes. We found that temperature changes induced substantial plasticity in terms of development time, body size and pre-adult survivorship, indicating that developmental temperature significantly affects growth and life history traits of C. megacephala. We also detected genetic differences among populations for body size and development time, and these two traits exhibited highly significant variations in the responses of different populations to various temperature conditions, indicating genetic differences among populations in terms of thermal reaction norms. The latitude of origin of the different populations (and hence mean temperature regimes in the environments from where the populations originated) did not appear to fully explain these genetic differences. In short, changes in development time and body size in C. megacephala can be regarded as adaptations to changing thermal regimes.     

Empirical evidence for fast temperature-dependent body size evolution in rotifers

Organisms tend to decrease in size with increasing temperature by phenotypic plasticity (the temperature-size rule; ectotherms) and/or genetically (Bergmann’s rule; all organisms). In this study, the evolutionary response of body size to temperature was examined in the cyclically parthenogenetic rotifer Brachionus plicatilis. Our aim was to investigate whether this species, already known to decrease in size with increasing temperature by phenotypic plasticity, presents a similar pattern at the genetic level. We exposed a multiclonal mixture of B. plicatilis to experimental evolution at low and high temperature and monitored body size weekly. Within a month, we observed a smaller size at higher temperature, as compared to body size at lower temperature. The pattern was consistent for the size of both mature females and eggs; rotifers kept at high temperature evolved to be on average 14% (after 2 weeks) and 3% (after 3 weeks) smaller than the ones kept at low temperature (10 and 5% in the case of eggs, respectively). We therefore found that B. plicatilis is genetically programmed to adjust its body size-to-environmental temperature.     

Complex latitudinal variation in the morphology of the kleptoparasitic spider Argyrodes kumadai associated with host use and climatic conditions

We examined complex geographical patterns in the morphology of a kleptoparasitic spider, Argyrodes kumadai, across its distributional range in Japan. To disentangle biotic and abiotic factors underlying morphological variation, latitudinal trends were investigated in two traits, body size and relative leg length, across separate transition zones for host use and voltinism. Statistical analyses revealed complex sawtooth clines. Adult body size dramatically changed at the transition zones for host use and voltinism, and exhibited a latitudinal decline following the converse to Bergmann’s cline under the same host use and voltinism in both sexes. A similar pattern was observed for relative leg length in females but not in males. A genetic basis for a part of observed differences in morphology was supported by a common-garden experiment. Our data suggest that local adaptation to factors other than season length such as resource availability (here associated with host use) obscures underlying responses to latitude.     

Effects of male apparent length on female preference for absolute body size in <Emphasis Type="Italic">Xiphophorus helleri</Emphasis>

Female green swordtails (Xiphophorus helleri) prefer males of longer body and sword lengths, which may reflect perceptual constraints. Larger males project a larger image onto the female’s retina at a given viewing distance, which may elicit greater stimulation of the visual system and a stronger behavioral response. This study investigates the effects of male apparent size, as affected by proximity and sword length, on female preference for absolute body size in X. helleri. Females were presented pairs of dummy males in ten treatments organized into six categories: (1) the Actual Length Control where identically sized males were held at different viewing distances from the female; (2) the Sword Compensation Treatment in which viewing distance and apparent length (body + sword) were held constant, pairing sworded with swordless dummies of equal total length; (3) the Body Length Preference Treatment where male distance from the female was held constant, while varying body size; (4) the Apparent (Sword) Length Preference Treatments where male distance from the female was held constant, while varying sword length; (5) the Apparent Length Controls where body length, sword length, and viewing distances varied while holding apparent length constant; and (6) the Experimental Treatment in which the smaller-sworded male of the pair appeared larger. In each of the Actual Length Control, Body, and Sword Length Treatments, females preferred the closer, larger-bodied, and longer-sworded males, respectively. However, females exhibited preferences in neither any of the Apparent Size Control treatments nor any of the Sword Compensation Treatment. Additionally, females in the Experimental Treatment preferred the apparently longer-sworded dummy to the male whose sword was actually longer. These results suggest that female preference for male body and sword length is based on the male’s apparent rather than actual size.     

Age and annual growth rate cause spatial variation in body size in Phrynocephalus przewalskii (Agamid)

Whether or not biogeographic rules dealing with spatial patterns of animal body sizes are valid for ectotherms is controversial. As the ectotherms grow all their lives, we explored the role of age and annual growth rate in body size variation in Phrynocephalus przewalskii in northern China. Morphological data were collected from 11 populations across a broad geographic gradient. Correlations between age, sex, climatic factors, and body size were analyzed using generalized linear model (GLM) and generalized linear mixed model (GLMM). GLM analysis indicated that the general body size of both sexes and the appendage size of females increased significantly with increasing temperature; however, the coefficient of determination was very small. GLMM analysis indicated that body size only correlated with age, whereas appendage size was affected by age, temperature, rainfall, and sunshine. Annual growth rates were positively correlated with temperature. We concluded that body size variation was mainly caused by age structure and plasticity of the growth rate in P. przewalskii and did not follow Bergmann''s rule; however, females followed Allen''s rule. Future studies to investigate the effect of energy restriction are needed to further understand the relationship between growth rate and body size. We also suggest that further studies on thermal advantage and sexual selection may be helpful to understand appendage size variation in P. przewalskii.     

Body surface temperature distribution in relation to body composition in obese women

Adipose tissue levels and human obesity are known to be associated with increased heat production. At the same time, subcutaneous adipose tissue provides an insulating layer that impedes heat loss. The energy implications of obesity and body thermoregulatory mechanisms remain relatively poorly understood. This study attempted to examine the potential relationship between body composition (subcutaneous and visceral fat) determined by bioimpedance as well as BMI (body mass index), and skin surface temperature distribution recorded at rest.One specific aim of this study was to draw a thermal map of body areas in obese women and compare this with women of normal body mass, and thus to identify body regions within which heat transfer is particularly impeded. As high fat content is a good insulator, it could reduce the body‘s ability to respond effectively to changes in environmental temperature, which would be problematic for thermal homeostasis. Our results showed that core temperature did not differ between obese and normal body mass participants, while skin temperature of most body surfaces was lower in obese subjects.The results of regression analysis showed that the mean body surface temperature (T_mean) decreased with increasing percentage of body fat (PBF) of the abdominal area. The opposite relationship was observed for the front area of the hand (simultaneous increase in T_mean and PBF). We also found a negative correlation between BMI and T_mean of the thigh areas, both the front and the back. From this it could be concluded that the mean body surface temperature is dependent on body fat.     

Ontogenetic and sexual differences of thermal biology and locomotor performance in a lacertid lizard,Eremias multiocellata

A viviparous lizard, Eremias multiocellata, was used to investigate the possible sexual and ontogenetic effects on selected body temperature, thermal tolerance range and the thermal dependence of locomotor performance. We show that adults are sexually dimorphic and males have larger bodies and heads than females. Adults selected higher body temperatures (34.5 vs. 32.4 °C) and could tolerate a broader range of body temperatures (8.1–46.8 vs. 9.1–43.1 °C) than juveniles. The sprint speed and maximum sprint distance increased with temperature from 21 °C to 33 °C, but decreased at 36 °C and 39 °C in both juveniles and adults. Adults ran faster and longer than juveniles at each tested temperature. Adult locomotor performance was not correlated with snout–vent length (SVL) or sex, and sprint speed was positively correlated with hindlimb length. Juvenile locomotor performance was positively correlated with both SVL and hindlimb length. The ontogenetic variation in selected body temperature, thermal tolerance and locomotor performance in E. multiocellata suggests that the effects of morphology on temperature selection and locomotor performance vary at different ontogenetic stages.     

The intrauterine origins of Hodgkin's lymphoma

Background: Both small and large body size at birth are now known to predict a range of chronic disorders in adult life, including certain cancers. These associations are thought to reflect “fetal programming.” This may lead to impairment of a small number of key systems including the immune system. Hodgkin's lymphoma is a disease of the immune system. We have therefore examined the association between Hodgkin's lymphoma and early development. Our hypothesis was that the disease would be associated with markers of poor fetal growth, specifically small body size or small placental size at birth. Methods: Using the Finnish Cancer Registry we identified patients with Hodgkin's disease in a cohort of 20,431 people born in Helsinki during 1924–44. Each person has a detailed birth record. Results: There were 12 patients with Hodgkin's disease, giving an incidence comparable to international rates. The disease was associated with prolonged gestation. For every additional week of gestation the hazard ratio was 1.37 (95% CI 1.00–1.87; p = 0.05). The disease was also associated with a short placental surface. After allowing for gestation, for each centimetre increase in surface length, the hazard ratio was 0.70 (0.53–0.92; p = 0.01). The disease was not associated with birth weight or length or maternal body size. Conclusions: We have shown that Hodgkin's lymphoma is associated with prolonged gestation and a short length of the placental surface. We speculate that Hodgkin's lymphoma is initiated by two events in fetal life. One, which is an immune event, is associated with prolonged gestation, while the other is associated with growth faltering.     

Thermoconformity strategy in the freshwater turtle Hydromedusa tectifera (Testudines,Chelidae) in its southern distribution area

Ectotherm species are not capable of generating metabolic heat; therefore, they present different strategies for regulating their body temperatures, ranging from a precise degree of thermoregulation to a passive thermoconformity with ambient temperatures. In reptiles, aerial basking is the most common mechanism for gaining heat. However, among aquatic reptiles, such as freshwater turtles, aquatic basking is also frequent. Hydromedusa tectifera is a turtle of exclusively aquatic and nocturnal habits widely distributed in South America. We studied the relationship between body temperature (Tb) of H. tectifera and its habitat, and explored the effects of sex, life stage and body size and mass on Tb. Fieldwork was conducted in two streams of a mountain area of central Argentina. We recorded cloacal temperature, size and mass of 84 turtles. We also determined individuals’ sex and life stage (adult/juvenile). Regarding ambient temperatures, we measured water temperature on the surface (Tsurf) and at depth of turtle capture (Tdepth) and air temperature. Mean Tb was 18.58 °C (Min = 10.20 °C; Max = 25.70 °C). Tsurf and Tdepth were highly correlated. Multi-model analysis using Akaike criterion indicated that Tb was strongly associated with water temperature, whereas air temperature and body size and mass did not show a significant effect. There was also no effect of turtle sex or life stage on Tb. Our results indicate that H. tectifera is a thermoconformer and eurythermal species. A nocturnal pattern of activity and a fully aquatic lifestyle are suggested as determinant factors.     

Changes in back fat thickness during late gestation predict colostrum yield in sows

Directing protein and energy sources towards lactation is crucial to optimise milk production in sows but how this influences colostrum yield (CY) remains unknown. The aim of this study was to identify associations between CY and the sow’s use of nutrient resources. We included 37 sows in the study that were all housed, fed and managed similarly. Parity, back fat change (ΔBF), CY and performance parameters were measured. We obtained sow serum samples 3 to 4 days before farrowing and at D1 of lactation following overnight fasting. These were analysed for non-esterified fatty acids (NEFA), urea, creatinine, (iso)butyrylcarnitine (C4) and immunoglobulins G (IgG) and A (IgA). The colostrum samples collected 3, 6 and 24 h after the birth of the first piglet were analysed for their nutrient and immunoglobulins content. The technical parameters associated with CY were parity group (a; parities 1 to 3=value 0 v. parities 4 to 7=value 1) and ΔBF D85-D109 of gestation (mm) (b): CY (g)=4290–842a–113b. (R²=0.41, P<0.001). The gestation length (P<0.001) and the ΔBF between D109 and D1 of lactation (P=0.050) were identified as possible underlying factors of the parity group. The metabolic parameters associated with CY were C4 at 3 to 4 days before farrowing (a), and 10logC4 (b) and 10logNEFA (c) at D1 of lactation: CY (g)=3582–1604a+1007b−922c (R²=0.39, P=0.001). The colostrum composition was independent of CY. The negative association between CY and ΔBF D85-D109 of gestation could not be further explained based on our data. Sows that were catabolic 1 week prior to farrowing seemed unable to produce colostrum to their full potential. This was especially the case for sows with parities 4 to 7, although they had a similar feed intake, litter birth weight and colostrum composition compared with parities 1 to 3 sows. In conclusion, this study showed that parity and the use of body fat and protein reserves during late gestation were associated with CY, indicating that proper management of the sow’s body condition during late gestation could optimise the intrinsic capacity of the sow’s CY.     

Bergmann's rule in amphibians: combining demographic and ecological parameters to explain body size variation among populations in the common toad Bufo bufo

Large‐scale patterns of body size variation are described by well‐known generalizations such as Bergmann’s rule; the generality and underlying causes of these patterns have been much debated. Intraspecific extension of this rule was tested in various ectotherms, and evidence was found for both Bergmann and converse Bergmann clines. In this study, we explored spatial patterns of variation in a widespread amphibian, the Common toad (Bufo bufo), along a 2240 km latitudinal gradient across Europe. We tested for covariation of adult body size, age and growth parameters with latitude, altitude, length of activity period and mean temperature during this period using both original and literature data. We selected 13 European populations, representing a latitudinal range from 43 to 63°N and altitudinal range from 15 to 1850 m a.s.l. The length of activity period (12–33 weeks) and T_mean (6.6–15.6°C) significantly decreased as latitude and altitude of these populations increased. Mean body size decreased as latitude increased (not with altitude), and increased with T_mean (not with length of activity period). Mean and minimal adult age increased with latitude and altitude, longevity increased with altitude only. Age increased as length of activity period decreased (not with T_mean). The growth coefficient (0.32–0.92 in males, 0.18–0.74 in females, available for six populations) decreased as altitude increased, and increased as both length of activity period and T_mean increased; latitudinal trend was non‐significant. Our analysis shows that B. bufo clearly exhibited a converse Bergmann cline along latitudinal gradient, but not along altitudinal gradient; the main effect of elevation was on age. The effects of ecological conditions also differed: body size increased with T_mean, while age parameters were related to the length of activity period. This study highlights that, to identify causal factors underlying general ecogeographical rules, we have to take into account different phases of the life cycle, co‐variation among life history traits and ecological factors acting on each of these traits. In amphibians with complex life cycles, lack of appropriate demographical or ecological data may affect our understanding of the variety of observed body size patterns.