The regeneration of a protected Sonoran Desert cactus since 1800 A.D. over 50,000 km2 of its range

The tall, branched saguaro cactus (Carnegiea gigantea) is perhaps the best-recognized symbol of the desert. However, little is known about the regeneration and future of the species outside of a few well-studied populations. In 2000, data for 537 saguaros were collected in 30 populations throughout the northern Sonoran Desert. A recently developed technique now provides a mechanism by which to reconstruct individual age at multiple populations. This new technique is based on a general growth curve with a site-specific adjustment factor, which I calculate based on local growth data and a recognized relationship with summer rain. Thus, the year of establishment was estimated for all saguaros in each of the populations individually, followed by the merging of all individuals, to create a single regeneration and survivorship curve for the combined regional dataset. Unlike other studies that only look at regeneration at one site, this is the first study to look at the long-term regeneration of the species over an area of more than 50,000 km², nearly the US portion of its range. The results suggest that over the long term, the population is quite stable, with a favorable period of regeneration in the late 1800s and early 1900s. It is also encouraging that the frequency of individuals that established in the most recent time period is relatively high. However, whether these youngest individuals will persist over the long-term in the face of future extreme freezing events (which can substantially thin populations) is not clear.     

Whisker growth in wild Eurasian badgers Meles meles: implications for stable isotope and bait marking studies

The use of biomarkers such as stable isotopes to study the foraging ecology and movement of animals is a rapidly expanding area of research. With respect to mammals, the analysis of inert keratinous tissue such as whiskers (vibrissae) is particularly attractive as they can be sequentially sampled to provide a long-term time series of individual movement or diet. However, in order to interpret data from such tissues researchers require details of growth rates and patterns, and also how these vary within populations. In this study, we use the fluorescent biomarker Rhodamine B to measure vibrissa growth rate and patterns in a wild population of Eurasian badgers. In addition, we compare stable isotope ratio values of blood and vibrissae in order to test whether vibrissae are retained for long periods following growth. We found that badger vibrissae grow at an average rate of 0.43 mm?day^?1 (range 0.23–0.83) such that single vibrissae sampled for stable isotope analysis contain an average of 104 days of ecological data. Age, sex and body condition did not affect growth rate, and there was no evidence of consistent individual differences in growth rate or long-term retention of vibrissae following growth. However, variation in growth rate within the population suggest that the temporal scales reflected in vibrissae may vary both between and within individuals, such that results are not always directly, temporally comparable. This research provides useful information for any future research using vibrissae in combination with biomarkers to study mammalian ecology.     

The allometry of saguaro height

The allometry of plant height H with respect to mean stem diameter D was determined based on 118 saguaro plants. The slope obtained for the reduced major axis regression analysis of the data was 2.36 ± 0.085, indicating that taller plants are disproportionately more slender than their shorter, presumably younger counterparts. The consequences of this positive, extremely anisometric relation on the elastic stability of stems were estimated by computing the critical buckling height H_crit for each of the 118 stems on the basis of the mean density-specific stiffness (i.e., the quotient of Young's elastic modulus E and bulk tissue density ρ) determined for a single section from a mature saguaro stem. E/ρ was nearly equivalent to that of tissue samples of sclerenchyma isolated from other plant species. Since the slope of H_crit vs. D equals ≈ 0.67 when E/ρ ≈ a constant, the safety-factor for saguaro stems (i.e., H_crit/H) appeared to be size-dependent such that it decreased with increasing plant height (i.e., H_crit/H ≈ D^-1.65). However, the mean safety-factor computed for the 118 saguaro specimens was 9.64, indicating that, on the average, plant height was well below H_crit. Additionally, circumstantial evidence suggests that saguaro stems become more stiff as they increase in size (and age) and that the rate of stem growth decelerates over time. The former would obtain a near size-independent safety-factor against elastic buckling while the latter protracts the time required to reach the critical buckling height. Comparisons among the allometries of H and H_crit for saguaro, other cacti, nonwoody, and highly branched tree species indicated that saguaro size overlaps with the lower size-range of the largest known dicot and gymnosperm tree specimens likely as a consequence of the high E/ρ of mature saguaro stems.     

Revisiting Bergmann's rule for saguaros (Carnegiea gigantea (Engelm.) Britt. and Rose): stem diameter patterns over space

Aim The influence of winter temperatures and other climate variables are explored to determine which variables are associated with saguaro stem diameter and to determine if Bergmann's rule is applicable to saguaros. Location The northern Sonoran Desert in Arizona, USA. Methods Thirty saguaro populations were sampled (height, diameter, number of branches), and after adjustment for population height structure, mean relative thickness of saguaros was calculated for each plot (population). Fifty‐seven climate variables were calculated for the thirty populations. Regression was run to determine which variables (cold winter, hot summer and precipitation) best predict relative thickness. Previous studies have demonstrated a significant positive relationship between winter precipitation and saguaro branching ( Drezner, 2003 , Ecography, in press). To determine if relative thickness may be an artefact of branching (branches), partial correlation analysis was employed. Results Mean March precipitation best predicts relative thickness. When only winter temperature variables are considered, none are significantly related to relative thickness. Relative thickness is not an artefact of branches. Main conclusions (1) Rainfall, not temperature, best predicts saguaro stem thickness. In addition, despite the focus on summer rains in the literature, winter precipitation is the best predictor of thickness. (2) Bergmann's rule is not applicable to saguaro populations as has been previously suggested [e.g. Niering et al. (1963) Science 142 , 15], as thickness does not increase significantly with latitude. In addition, the suggested mechanism for Bergmann's rule, cold winter temperature, does not significantly predict saguaro stem diameters over the area studied. In some populations that experience high winter rainfall as well as cold temperatures, individuals likely derive thermal benefits from a larger stem diameter; however, the trend is not observed over the area studied and it does not appear to be adaptive.     

Floral biology of the saguaro (Cereus giganteus)

A saguaro cactus (Cereus giganteus) produces an average of 295 flowers per season, each of which produces 286 mg fresh weight of pollen and 543 mg of nectar containing 24% sugar. At 7600 pollen grains/mg pollen, the yearly output per saguaro plant is 6.4×10⁸ grains. Based on the measured saguaro density of 6.56 plants/ha, 553 g/ha of pollen is produced yearly. The enormous variation among individual plants in terms of flower numbers and floral bloom patterns is documented.Honey bees (Apis mellifera L.), the main collectors of saguaro pollen, collect an average of 12.2 mg pollen per foraging trip and can thus harvest 23.5 pollen loads from one flower. An average honey bee colony collects 290 g of saguaro pollen over the season, which is 24.4% of their total intake. Individual colonies exhibit wide variation in pollen collecting activities with some closely tracking the pollen resource and others almost totally ignoring it. The average for seven colonies indicates that even though variation is great the overall trend is toward closely tracking and exploiting the saguaro pollen resource. Based on the pollen productivity of saguaro and a hypothetical 90% pollen harvesting efficiency of bees, the pollen harvest potential of the saguaro environment is 1.72 colony equivalents of pollen/ha and 0.5/ha for saguaro alone. This is the first quantitative reporting of the total pollen productivity and pollen resource utilization for any plant and an opportunistic pollinator.     

Calculating the intrinsic growth rate: comparison of definition and model

It was shown that well known equation r = ln[N(t2)/N(t1)]/(t2 - t1) is the definition of the average value of intrinsic growth rate of population r within any given interval of time t2-t1 and changing arbitrarity its numbers N(t). The common opinion considering the equation as suitable only for exponentially growing population was found to be incorrect. The fundamentally different approach is based on the calculation of r within the framework of demographic model, realized as Euler - Lotka equation or population projection matrices. However this model requires simultaneous realization of several assumptions improbable for natural populations: exponential change in population size, stable age structure and maintaining constant age-dependent birth and death rates. The calculation of r by definition requires the data on the dynamics of population numbers, whereas calculation on the basis of the model requires the demographic tables of birth and death rate, but not the population numbers. With the example of American ginseng it was shown that evalution of r by definition and model approaches could produce opposite results.     

Probabilistic Gompertz model of irreversible growth

Characterizing organism growth within populations requires the application of well-studied individual size-at-age models, such as the deterministic Gompertz model, to populations of individuals whose characteristics, corresponding to model parameters, may be highly variable. A natural approach is to assign probability distributions to one or more model parameters. In some contexts, size-at-age data may be absent due to difficulties in ageing individuals, but size-increment data may instead be available (e.g., from tag-recapture experiments). A preliminary transformation to a size-increment model is then required. Gompertz models developed along the above lines have recently been applied to strongly heterogeneous abalone tag-recapture data. Although useful in modelling the early growth stages, these models yield size-increment distributions that allow negative growth, which is inappropriate in the case of mollusc shells and other accumulated biological structures (e.g., vertebrae) where growth is irreversible. Here we develop probabilistic Gompertz models where this difficulty is resolved by conditioning parameter distributions on size, allowing application to irreversible growth data. In the case of abalone growth, introduction of a growth-limiting biological length scale is then shown to yield realistic length-increment distributions.     

Variation in age and height of onset of reproduction in the saguaro cactus (<Emphasis Type="Italic">Carnegiea gigantea</Emphasis>) in the Sonoran Desert

The objective of this study is to determine the height and age at which reproduction begins (i.e., production of flowers and fruits; the transition to adulthood) in the giant saguaro cactus (Carnegiea gigantea) in four geographically and environmentally distinct populations, and to relate observed variability to environmental differences. The onset of reproduction has been estimated at a height of 2.2 m in near optimal conditions. This value has been widely accepted and applied to populations in less optimal conditions, although variations under less ideal conditions have not been investigated. In addition, previous research has demonstrated that Carnegiea growth rates are highly variable over their range. Thus, even if 2.2 m is a consistent transition height to adulthood over their range, the age of individuals in different populations would be different. I investigate the age and height at which this transition occurs. The author sampled the heights of the shortest reproductive individuals and the tallest non-reproductive individuals to estimate the mean height of the onset of flowering in each of four locales in the northern Sonoran Desert. Using a previously published age-height-growth model, the mean age of the start of reproduction was also computed for the four sites. ANOVA and t-tests were used to compare the average transition to adulthood across sites by both age and height. Statistical results are robust and significant variations in the onset of reproduction are observed by both age and height across the four sites. Saguaro National Park and Organ Pipe Cactus National Monument individuals are transitioning to adulthood, on average, at younger ages and shorter heights than the other two locales. At the arid and marginal Kofa site, individuals that established during the regeneration peak of the late 1800s-early 1900s are only now becoming reproductive (individuals that established around 1899), while at Saguaro National Park, on average, individuals that established in the 1950s are already transitioning to adulthood. These results have implications for regeneration, particularly in marginal locales where regeneration is already limited.     

Probability density functions useful for parametrization of heterogeneity in growth and allometry data

This paper addresses the problem of modelling heterogeneous individual characteristics in a population. A flexible unified approach for stochastic parametrization dynamics of the distribution in population data is proposed. To approximate data with multiple observations per individual, models based on Markov processes are constructed. The method can be applied to scalar or multivariate characteristics, and its application to growth and allometry data is considered. Different stochastic versions of known growth and allometry functions are developed, which enable wide applicability. Simple informative growth indices are calculated as the moments of distribution. The three-parameter Gompertz growth model for size-at-age data was reparametrized to a size-increment data model with two parameters. An erratum to this article is available at .     

Distribution and growth of brown trout in pristine headwaters of Central Europe

The majority of stream-dwelling salmonid populations in Europe are affected by artificial stocking and the fragmentation of riverine ecosystems. The present study was performed in the unique pristine headwaters of the Otava River in the Elbe catchment area of the Czech Republic. The aim was to investigate the spatial distribution and individual growth pattern of brown trout, Salmo trutta, populations. Twenty sites in two main streams and their tributaries were sampled twice a year (spring and autumn) during the period 2005–2011. The sampling sites were grouped into fourteen so-called synchronised population units within the boundaries of three populations, according to analyses of synchrony in population abundance. The individual growth of juveniles (age-0, age-1) varied between all three spatial units (sampling sites, synchronised population units and populations), while the individual growth of adults (age-2 and older) did not. The distinctiveness regarding individual growth and demographic independence among the synchronised population units and populations indicates their suitability for use as population units for management purposes.     

Challenges of DNA-Based Mark-Recapture Studies of American Black Bears

Abstract: We explored whether genetic sampling would be feasible to provide a region-wide population estimate for American black bears (Ursus americanus) in the southern Appalachians, USA. Specifically, we determined whether adequate capture probabilities (p > 0.20) and population estimates with a low coefficient of variation (CV < 20%) could be achieved given typical agency budget and personnel constraints. We extracted DNA from hair collected from baited barbed-wire enclosures sampled over a 10-week period on 2 study areas: a high-density black bear population in a portion of Great Smoky Mountains National Park and a lower density population on National Forest lands in North Carolina, South Carolina, and Georgia. We identified individual bears by their unique genotypes obtained from 9 microsatellite loci. We sampled 129 and 60 different bears in the National Park and National Forest study areas, respectively, and applied closed mark-recapture models to estimate population abundance. Capture probabilities and precision of the population estimates were acceptable only for sampling scenarios for which we pooled weekly sampling periods. We detected capture heterogeneity biases, probably because of inadequate spatial coverage by the hair-trapping grid. The logistical challenges of establishing and checking a sufficiently high density of hair traps make DNA-based estimates of black bears impractical for the southern Appalachian region. Alternatives are to estimate population size for smaller areas, estimate population growth rates or survival using mark-recapture methods, or use independent marking and recapturing techniques to reduce capture heterogeneity.     

Morphometrics and pattern of growth in wild sifakas (Propithecus edwardsi) at ranomafana national park,madagascar

We summarize morphometric data collected over a period of 22 years from a natural population of rainforest sifakas (Propithecus edwardsi) at Ranomafana National Park, Madagascar, and we use those data to document patterns of growth and development. Individually identified, known‐age sifakas were successfully captured, measured, and released. We found that body segment lengths increased faster during growth than did body mass, with individuals attaining adult lengths earlier than adult mass. Females can begin reproducing before they are fully grown, but this may not be common. With the exception of hand length, we found no significant sex difference in any adult metric including body mass, chest, and limb circumferences, body segment lengths, and canine tooth height; however, body masses of individual females fluctuated more, independently of pregnancy, than did those of males. We found considerable interannual fluctuation in body mass with single individuals differing more within the same season in different years than from season to season in the same year. Such body mass fluctuation may be a consequence of eastern Madagascar's variable and unpredictable environment in which rainfall during any selected month varies from year to year. Am. J. Primatol. 73:155–172, 2011. © 2010 Wiley‐Liss, Inc.     

A Rapid Method to Estimate Population Variables for African Elephants

Abstract: We developed a noninvasive method to estimate reproductive and survival parameters for free-ranging African savannah elephants (Loxodonta africana africana) and used these to estimate finite population growth rates. We used published data from 2 populations with known growth rates and birth and survival histories to validate our technique. Based on body measurements, our method yielded estimates of age at first and last calving, calving interval, and age-specific survival rates that were similar to those determined during long-term studies at both Addo Elephant National Park and Amboseli National Park. Our technique generated population data required to estimate population growth rates. The method may be particularly useful where censuses yield imprecise estimates or where long-term population data are unavailable. (JOURNAL OF WILDLIFE MANAGEMENT 72(3):822–829; 2008)     

Molecular evolution and population genetics of two Drosophila mettleri cytochrome P450 genes involved in host plant utilization

Understanding the genetic basis of adaptation is one of the primary goals of evolutionary biology. The evolution of xenobiotic resistance in insects has proven to be an especially suitable arena for studying the genetics of adaptation, and resistant phenotypes are known to result from both coding and regulatory changes. In this study, we examine the evolutionary history and population genetics of two Drosophila mettleri cytochrome P450 genes that are putatively involved in the detoxification of alkaloids present in two of its cactus hosts: saguaro (Carnegiea gigantea) and senita (Lophocereus schottii). Previous studies demonstrated that Cyp28A1 was highly up-regulated following exposure to rotting senita tissue while Cyp4D10 was highly up-regulated following exposure to rotting saguaro tissue. Here, we show that a subset of sites in Cyp28A1 experienced adaptive evolution specifically in the D. mettleri lineage. Moreover, neutrality tests in several populations were also consistent with a history of selection on Cyp28A1. In contrast, we did not find evidence for positive selection on Cyp4D10, although this certainly does not preclude its involvement in host plant use. A surprising result that emerged from our population genetic analyses was the presence of significant genetic differentiation between flies collected from different host plant species (saguaro and senita) at Organ Pipe National Monument, Arizona, USA. This preliminary evidence suggests that D. mettleri may have evolved into distinctive host races that specialize on different hosts, a possibility that warrants further investigation.     

Heparin-binding growth factor/prostatropin attenuates inhibition of rat prostate tumor epithelial cell growth by transforming growth factor type beta

Summary Normal rat prostate epithelial cell growth requires both epidermal growth factor and heparin-binding growth factor/prostatropin. In contrast, epithelial cells derived from the transplantable Dunning R3327H rat tumor require either epidermal growth factor or heparin-binding growth factor/prostatropin. Transforming growth factor type beta inhibited normal epithelial cell growth. Transforming growth factor beta inhibited epidermal growth factor-dependent growth of tumor epithelial cells, independent of epidermal growth factor concentrations. Transforming growth factor beta increased the effective dose of heparin-binding growth factor type 1 required to support tumor epithelial cell growth by 10-fold but saturating levels of heparin-binding growth factor type 1 (290 pM) completely attenuated the inhibitory effect of transforming growth factor beta. These results suggest that prostate tumor epithelial cells may escape the inhibitory effect of transforming growth factor beta as a consequence of alteration of the concurrent requirement for both epidermal growth factor (or homologues) and heparin-binding growth factors. This work was supported by NCI Grant CA37589. Editor’s Statement The observation that heparin-binding growth factor/prostatropin can counteract the inhibitory effect of transforming growth factor beta in prostate epithelial cells may help explain how some cancers avoid the action of growth inhibitors and provides a model for studying how inhibitory peptides overcome the stimulatory signals generated by growth factors.     

Relationships between basic life history parameters and population size

We consider an ideal population with a stable age composition changing according Lotka equation. Additional assumptions are made concerning the constancy of population size, independence of specific mortality rate on age, and linear dependence of female fecundity on its weight. A relationship has been obtained [formula: see text] where N0 is initial numbers of a generation, N[alpha, omega] is total numbers of the mature part of the population, w[alpha, omega] is a mean weight of a mature individual, s is sex ratio, c is specific fecundity (per unit of weight) and l0 is the probability of larval surviving. The growth of an individual is described by the Bertalanffy function. Methods of calculation of life history parameters are discussed. A method is proposed to calculate the age of maturity (alpha) and at the end (omega) of the reproduction period as first and second inflection points of the growth rate curve. Based upon data on development of 27 populations of several species of fishes of inland waters of Russia the following relationship have been obtained: [formula: see text] for populations with [formula: see text] < or = 100 g, [formula: see text] for populations with [formula: see text] > 100 g, and [formula: see text] for all populations.     

Comparison of growth curves of mice selected and unselected for postweaning gain

Summary Mice were sampled from a line selected for increased postweaning weight gain from three to six weeks and from a randombred control line originating from the same base population. Body weights were recorded at each of 14 ages from day 5 to day 98. The Richards and logistic growth functions were fitted to the growth trajectories of each individual mouse by a generalized non-linear least squares procedure. Estimated growth parameters (asymptotic weight, rate, shape of curve, age and weight at inflection, mean absolute growth rate and mean relative growth rate) were computed for each individual. The effects of line, litter within line, sex and line × sex interactions on these estimated parameters were then studied.Both the Richards and logistic functions fitted the data equally well and the plotted trajectories coincided over most of the growth curve. There was excellent agreement between the estimates of asymptotic weight and both age and weight at inflection based on the different functions. However, both functions apparently underestimated the asymptotic weight.Analyses of the line differences showed that selection for postweaning gain increased the mean absolute growth rate over the entire curve but had no effect on the relative growth rate or the shape of the growth curve. Full-sib analyses suggested the presence of considerable genetic variation and some high genetic correlations among the estimated growth parameters.Paper number 2942 of the Journal Series of the North Carolina State University Agricultural Experiment Station, Raleigh, North Carolina. This research was supported in part by Public Health Service Research Grant GM 11546. Computing services were supported by NIH Grant FR-00011.     

Estimating age at maturation and energy-based life-history traits from individual growth trajectories with nonlinear mixed-effects models

A new method is presented to estimate individuals’ (1) age at maturation, (2) energy acquisition rate, (3) energy expenditure for body maintenance, and (4) reproductive investment, and the multivariate distribution of these traits in a population. The method relies on adjusting a conceptual energy allocation model to individual growth curves using nonlinear mixed-effects modelling. The method’s performance was tested using simulated growth curves for a range of life-history types. Individual age at maturation, energy acquisition rate and the sum of maintenance and reproductive investment rates, and their multivariate distribution, were accurately estimated. For the estimation of maintenance and reproductive investment rates separately, biases were observed for life-histories with a large imbalance between these traits. For low reproductive investment rates and high maintenance rates, reproductive investment rate estimates were strongly biased whereas maintenance rate estimates were not, the reverse holding in the opposite situation. The method was applied to individual growth curves back-calculated from otoliths of North Sea plaice (Pleuronectes platessa) and from scales of Norwegian spring spawning herring (Clupea harengus). For plaice, maturity ogives derived from our individual estimates of age at maturation were almost identical to the maturity ogives based on gonad observation in catch samples. For herring, we observed 51.5 % of agreement between our individual estimates and those directly obtained from scale reading, with a difference lower than 1 year in 97 % of cases. We conclude that the method is a powerful tool to estimate the distribution of correlated life-history traits for any species for which individual growth curves are available.     

Clinical standards for growth in height of Belgian boys and girls,aged 2 to 18 years

The present paper presents the first clinical standard for growth in height of Belgian boys and girls, based on purely longitudinal data. Growth charts are provided with centiles of height for age along with growth curves of the typical early, average and late maturing child in the population. These new standards show the classical features of cross-sectional standards, but above that, they also provide information about the variability in individual growth patterns, as a result of variation in maturation. Average adult height is 176.6 cm (SD=6.3 cm) in boys and 163.3 cm (SD=5.7 cm) in girls. The representativity of these new standards with respect to the actual Belgian population has been by comparison with recent cross-sectional data, collected on a large number of subjects. These standards should be applied in all situations where interest lies in the evaluation of the normality of a child's growth pattern over some length of time and will therefore find their usefulness in clinical follow-up studies of growth.