Identification and quantification of projectile impact marks on bone: new experimental insights using osseous points.

Shifts in projectile technology potentially document human evolutionary milestones, such as adaptations for different environments and settlement dynamics. A relatively direct proxy for projectile technology is projectile impact marks (PIM) on archaeological bones. Increasing awareness and publication of experimental data sets have recently led to more identifications of PIM in various contexts, but diagnosing PIM from other types of bone-surface modifications, quantifying them, and inferring point size and material from the bone lesions need more substantiation. Here, we focus on PIM created by osseous projectiles, asking whether these could be effectively identified and separated from lithic-tipped weapons. We further discuss the basic question raised by recent PIM research in zooarchaeology: why PIM evidence is so rare in archaeofaunal assemblages (compared to other human-induced marks), even when they are explicitly sought. We present the experimental results of shooting two ungulate carcasses with bone and antler points, replicating those used in the early Upper Paleolithic of western Eurasia. Half of our hits resulted in PIM, confirming that this modification may have been originally abundant. However, we found that the probability of a skeletal element to be modified with PIM negatively correlates with its preservation potential, and that much of the produced bone damage would not be identifiable in a typical Paleolithic faunal assemblage. This quantification problem still leaves room for an insightful qualitative study of PIM. We complement previous research in presenting several diagnostic marks that retain preservation potential and may be used to suggest osseous, rather than lithic, projectile technology. Supplementary Information: The online version contains supplementary material available at 10.1007/s12520-024-01944-3.

Does Health & Her app use improve menopausal symptoms? A longitudinal cohort study.

OBJECTIVES: The Health & Her app provides menopausal women with a means of monitoring their symptoms, symptom triggers and menstrual periods, and enables them to engage in a variety of digital activities designed to promote well-being. This study aimed to examine whether sustained weekly engagement with the app is associated with improvements in menopausal symptoms. DESIGN: A pre-post longitudinal cohort study. SETTING: Analysed data collected from Health & Her app users. PARTICIPANTS: 1900 women who provided symptom data via the app across a 2-month period. PRIMARY AND SECONDARY OUTCOME MEASURES: Symptom changes from baseline to 2 months was the outcome measure. A linear mixed effects model explored whether levels of weekly app engagement influenced symptom changes. Secondary analyses explored whether app-usage factors such as total number of days spent logging symptoms, reporting triggers, reporting menstrual periods and using in-app activities were independently predictive of symptom changes from baseline. Covariates included hormone replacement therapy use, hormonal contraceptive use, present comorbidities, age and dietary supplement use. RESULTS: Findings demonstrated that greater engagement with the Health & Her app for 2 months was associated with greater reductions in symptoms over time. Daily use of in-app activities and logging symptoms and menstrual periods were each independently associated with symptom reductions. CONCLUSIONS: This study demonstrated that greater weekly engagement with the app was associated with greater reductions in symptoms. It is recommended that women be made aware of menopause-specific apps, such as that provided by Health & Her, to support them to manage their symptoms.

Symptom monitoring improves physical and emotional outcomes during menopause: a randomized controlled trial.

ABSTRACT: Objectives: A recent systematic review suggested that symptom monitoring can result in reductions in menopausal symptoms and improvements in health-related behaviors. To date, no studies have experimentally investigated whether symptom monitoring could be a beneficial intervention during the menopause transition.Methods: One hundred perimenopausal and postmenopausal women (mean age, 46 y; SD, 8 y) were randomized into either a monitoring-intervention or control group. A mixed between/within design was used, with group membership (ie, monitoring-intervention or control) as the between-subjects component and time (ie, baseline and 2-wk follow-up) as the within-subjects component. Dependent variables included symptom reductions and emotional reactions as measured via the Daily Record Keeping form. Secondary outcomes included help-seeking, communication, medical decision making, health awareness, self-efficacy, and health anxiety.Results: A linear mixed-effects model demonstrated that the monitoring-intervention group reported a 42% reduction in physical symptoms at follow-up versus a 12% reduction in the control group: ρ = 0.009, ß = 6.3, 95% CI (1.5-11). Negative emotions also significantly reduced in the monitoring-intervention group but did not alter in the control group: ρ < 0.001, ß = 3.4, and 95% CI (1.6-5.2). These effects remained significant after controlling for potential moderator variables such as trait neuroticism and coping preferences and potential confounders such as medical and demographic characteristics. Variances in other health outcomes were nonsignificant.Conclusions: Findings demonstrated that symptom monitoring reduced symptoms and negative emotions within a perimenopausal and postmenopausal sample, and these outcomes endured after controlling for key moderators and covariates. However, symptom monitoring was not related to improvements in health-related behavioral outcomes, which contrasts with previous findings. These findings show that symptom monitoring may be useful within healthcare settings by providing perimenopausal and postmenopausal women with a simple and accessible means of symptom alleviation while they await treatment or medical consultation.

Evaluating the Effects of Symptom Monitoring on Menopausal Health Outcomes: A Systematic Review and Meta-Analysis.

Evidence suggests that monitoring and appraising symptoms can result in increased engagement in medical help-seeking, improved patient-doctor communication, and reductions in symptom prevalence and severity. To date, no systematic reviews have investigated whether symptom monitoring could be a useful intervention for menopausal women. This review explored whether symptom monitoring could improve menopausal symptoms and facilitate health-related behaviours. Results suggested that symptom monitoring was related to improvements in menopausal symptoms, patient-doctor communication and medical decision-making, heightened health awareness, and stronger engagement in setting treatment goals. Meta-analyses indicated large effects for the prolonged use of symptom diaries on hot flush frequencies. Between April 2019 and April 2021, PsychInfo, EMBASE, MEDLINE, CINAHL, Cochrane, ProQuest, PsychArticles, Scopus, and Web of Science were searched. Eighteen studies met the eligibility criteria and contributed data from 1,718 participants. Included studies quantitatively or qualitatively measured the impact of symptom monitoring on menopausal populations and symptoms. Research was narratively synthesised using thematic methods, 3 studies were examined via meta-analysis. Key themes suggest that symptom monitoring is related to improvements in menopausal symptoms, improved patient-doctor communication and medical decision-making, increased health awareness, and stronger engagement in goal-setting behaviours. Meta-analysis results indicated large effects for the prolonged use of symptom diaries on hot flush frequency: 0.73 [0.57, 0.90]. This review is limited due to the low number of studies eligible for inclusion, many of which lacked methodological quality. These results indicate that symptom monitoring has potential as an effective health intervention for women with menopausal symptoms. This intervention may be beneficial within healthcare settings, in order to improve patient-doctor relations and adherence to treatment regimes. However, findings are preliminary and quality assessments suggest high risk of bias. Thus, further research is needed to support these promising outcomes. Systematic Review Registration Number: https://www.crd.york.ac.uk/prospero/display_record.php?, PROSPERO, identifier: CRD42019146270.

Adaptive variation in the development of extraembryonic membranes of gekkotan embryos: A meta-analytical approach.

Highly mineralized rigid-shelled eggs characterize one lineage of gekkotans. In contrast, poorly mineralized flexible-shelled eggs characterize basal lineages of gekkotans and all other squamates. Low oxygen permeability of rigid-shelled eggs is associated with small eggs and hatchlings, and long incubation lengths compared to flexible-shelled gekkotan eggs. These features represent a demographic cost for species with rigid-shelled eggs. This cost is offset, at least in part, because mortality due to desiccation and predation is reduced for rigid-shelled eggs relative to flexible-shelled eggs. Developmental traits may also compensate for the low oxygen permeability of rigid-shelled eggs. Oviposition, for example, occurs at earlier developmental stages for gekkotans with rigid- versus flexible-shelled eggs. Such early oviposition facilitates development because eggs move from the relatively hypoxic oviduct to the much better oxygenated nest environment. In this study, I tested the hypothesis that the growth of the yolk sac (YS) and chorioallantoic membrane (CAM) of gekkotans with rigid-shelled eggs is initiated and completed earlier than those of gekkotans with flexible-shelled eggs. I measured the surface area of eggs covered by the YS and CAM from oviposition to hatching and determined which of four nonlinear models provide the best fit for growth curves. I also compiled a data set on embryonic metabolism of gekkotans and other lizards in order to place growth of the YS and CAM in the context of energy utilization of lizard embryos overall. Growth of the YS and CAM of gekkotans with rigid-shelled eggs is accelerated relative to that of gekkotans with flexible-shelled eggs and may serve to separate the cost of YS and CAM development from that of the embryo itself. Adaptive variation in YS and CAM development may extend to birds, crocodilians, and turtles as they also exhibit life history variation that affects oxygen availability to embryos during development.

Agricultural land use creates evolutionary traps for nesting turtles and is exacerbated by mercury pollution.

The rate of anthropogenic habitat conversion often exceeds the rate of natural ecological and evolutionary processes, which sometimes creates mismatches between environmental cues and adaptive behaviors. In numerous species of aquatic turtles, nest site selection is primarily based on cues related to high solar exposure, which leads females to select sites where humans have disturbed habitat. These disturbed sites are often contaminated by pollutants, such as mercury. Despite the fact that anthropogenic disturbances often co-occur, few studies have examined the interactive influence of major global changes on animal development. Using Chelydra serpentina, we investigated the individual and interactive effects of crop agriculture and mercury pollution on hatch success and offspring phenotype. We hypothesized that following nesting, rapid crop growth would shade and cool nests in agricultural fields and subsequently negatively impact embryonic development. Agricultural and control nests were similar in temperature at the time of oviposition, but temperatures diverged as crops grew: agricultural nests averaged 2.5°C cooler than control nests over the course of incubation. In laboratory and field experiments, we found that turtles incubated under agricultural thermal regimens took longer to hatch, hatched at smaller body sizes, lost more mass, and had lower posthatching growth rates. Additionally, thermal conditions associated with agricultural land use interacted with mercury contamination to decrease hatching success. To our knowledge, this is the first documentation of negative interactive effects of mercury pollution and habitat quality on early vertebrate development and highlights the importance of examining the combined influence of anthropogenic global changes on organisms.

Earthworms (Oligochaeta: Lumbricidae) of Interior Alaska.

Earthworms in the family Lumbricidae in Alaska, which are known from coastal regions, primarily in south-central and south-eastern Alaska, are thought to be entirely non-native and have been shown to negatively impact previously earthworm-free ecosystems in study regions outside of Alaska. Despite occasional collections by curious citizens, there had not been a standardised earthworm survey performed in Interior Alaska and no published records exist of earthworms species from this region. Mustard extraction was used to sample six locations that differed in elevation, mostly in the College region of Fairbanks, Alaska. Two of the six locations yielded earthworms. There was no relationship between earthworm abundance and elevation (p = 0.087), although our sample size was small. Our sampling, combined with specimens in the University of Alaska Museum, has documented four exotic species and one presumed native species of lumbricid earthworms in Interior Alaska.

Developmental plasticity in reptiles: Insights into thermal and maternal effects on chameleon phenotypes.

Embryonic environments affect a range of phenotypic traits including sex and reproductive success. I determined (1) how the interaction between incubation temperature and egg size affects sex allocation of Chamaeleo calyptratus and (2) how incubation temperature and maternal parent (clutch) affect water uptake by eggs and body size, growth, and climbing speed of hatchlings and juveniles. Eggs from five clutches were exposed to five temperature treatments with clutches replicated within and among treatments. Temperature affected sex, but only when egg size was included as a factor in analyses. At intermediate (28°C) temperatures, daughters were more likely to be produced from large eggs and sons more likely to be produced from small eggs, while at 25 and 30°C, the pattern of sex allocation was reversed. Temperature and clutch affected water uptake and body size. Nonetheless, the direction of temperature and clutch effects on water uptake by eggs and on the size of hatchlings were not the same and the direction of temperature effects on body sizes of hatchlings and juveniles differed as well. Clutch affected hatchling size but not juvenile size and growth rate. Clutch, but not incubation temperature, affected climbing speed, but the fastest hatchlings were not from the same clutches as the fastest juveniles. The independent effects of incubation temperature and clutch indicate that hatchling phenotypes are influenced largely by conditions experienced during incubation, while juvenile phenotypes are influenced largely by conditions experienced in the rearing environment.

Developmental origin of limb size variation in lizards.

In many respects, reptile hatchlings are fully functional, albeit miniature, adults. This means that the adult morphology must emerge during embryonic development. This insight emphasizes the connection between the mechanisms that generate phenotypic variation during embryonic development and the action of selection on post-hatching individuals. To determine when species-specific differences in limb and tail lengths emerge during embryonic development, we compared allometric patterns of early limb growth of four distantly related species of lizards. The major questions addressed were whether early embryonic limb and tail growth is characterized by the gradual (continuous allometry) or by the abrupt emergence (transpositional allometry) of size differences among species. Our observations supported transpositional allometry of both limbs and tails. Species-specific differences in limb and tail length were exhibited when limb and tail buds first protruded from the body wall. Genes known to be associated with early limb development of tetrapods are obvious targets for studies on the genetic mechanisms that determine interspecific differences in relative limb length. Broadly comparative studies of gene regulation would facilitate understanding of the mechanisms underlying adaptive variation in limb size, including limb reduction and loss, of squamate reptiles.

Rigid shells enhance survival of gekkotan eggs.

The majority of lizards and snakes produce permeable parchment-shelled eggs that require high moisture conditions for successful embryonic development. One clade of gekkotan lizards is an exception; females produce relatively impermeable rigid-shelled eggs that normally incubate successfully under low moisture conditions. I tested the hypothesis that the rigid-shell increases egg survival during incubation, but only under low moisture conditions. To test this hypothesis, I incubated rigid-shelled eggs of Chondrodactylus turneri under low and under high moisture conditions. Eggs were incubated with parchment-shelled eggs of Eublepharis macularius to insure that incubation conditions were suitable for parchment-shelled eggs. Chondrodactylus turneri eggs had very high survival (>90%) when they were incubated under low moisture conditions. In contrast, eggs incubated under high moisture conditions had low survival overall, and lower survival than those of the parchment-shelled eggs of E. macularius. Mortality of C. turneri and E. macularius eggs incubated under high moisture conditions was the result of fungal infection, a common source of egg mortality for squamates under laboratory and field conditions. These observations document high survival of rigid-shelled eggs under low moisture conditions because eggs escape from fungal infection. Highly mineralized rigid shells also make egg survival independent of moisture availability and may also provide protection from small invertebrates in nature. Enhanced egg survival could thus compensate for the low reproductive output of gekkotans that produce rigid-shelled eggs.

Long-term data reveal a population decline of the tropical lizard Anolis apletophallus, and a negative affect of el nino years on population growth rate.

Climate change threatens biodiversity worldwide, however predicting how particular species will respond is difficult because climate varies spatially, complex factors regulate population abundance, and species vary in their susceptibility to climate change. Studies need to incorporate these factors with long-term data in order to link climate change to population abundance. We used 40 years of lizard abundance data and local climate data from Barro Colorado Island to ask how climate, total lizard abundance and cohort-specific abundance have changed over time, and how total and cohort-specific abundance relate to climate variables including those predicted to make the species vulnerable to climate change (i.e. temperatures exceeding preferred body temperature). We documented a decrease in lizard abundance over the last 40 years, and changes in the local climate. Population growth rate was related to the previous years' southern oscillation index; increasing following cooler-wetter, la niña years, decreasing following warmer-drier, el nino years. Within-year recruitment was negatively related to rainfall and minimum temperature. This study simultaneously identified climatic factors driving long-term population fluctuations and climate variables influencing short-term annual recruitment, both of which may be contributing to the population decline and influence the population's future persistence.

Developmental sequences of squamate reptiles are taxon specific.

Recent studies in comparative vertebrate embryology have focused on two related questions. One concerns the existence of a phylotypic period, or indeed any period, during development in which sequence variation among taxa is constrained. The second question concerns the degree to which developmental characters exhibit a phylogenetic signal. These questions are important because they underpin attempts to understand the evolution of developmental characters and their links to adult morphology. To address these questions, we compared the sequence of developmental events spanning the so-called phylotypic period of vertebrate development in squamate reptiles (lizards and snakes), from the formation of the primary optic placode to the first appearance of scale anlagen. We used Bayesian phylogenetic ancestral state reconstruction analyses and estimates of Bayesian posterior probabilities of the rank order of developmental events to determine the level of support for phylogenetically associated variation in development. We assessed the amount of variation in event sequences by plotting the proportions of reconstructed ranks (excluding unlikely events, PP < 0.05) associated with each event. Sequence variability was the lowest towards the middle of the phylotypic period and involved three events (allantois contacts chorion, maximum number of pharyngeal slits, and appearance of the apical epidermal ridge [AER]); these events each had only two reconstructed ranks. Squamate clades also differed in the rank order of developmental events. Of the 20 events in our analyses, 12 had strongly supported (PP ≥ 0.95) sequence ranks that differed at two or more internal nodes of the tree. For example, gekkotans are distinguished by the late appearance of the allantois bud compared to all other squamates (ranks 7 and 8 vs. rank 3, respectively) and Serpentes are distinguished by the earlier completion of torsion (rank 3) compared to acrodonts and pleurodonts (ranks 7 and 5, respectively). Clade specific sequences of developmental events mean that investigators should not extend observations on the development on particular squamate species to distantly related taxa for use in comparative studies.

Does low gas permeability of rigid-shelled gekkotan eggs affect embryonic development?

Parchment-shelled eggs are characteristic of most squamates, including the basal clades of gekkotan lizards. The majority of gekkotan lizards, however, produce rigid-shelled eggs that are highly impermeable to gas exchange; eggs are laid in dry sites and experience a net loss of water during incubation. We tested the hypothesis that the 1,000-fold lower rate of oxygen diffusion through the shells of rigid- compared to parchment-shelled eggs imposes a physiological cost on development. To do this, we contrasted species with rigid and with parchment shells with regards to (1) rates of embryonic metabolism and (2) rates and patterns of development of the yolk sac and chorioallantois, the vascularized extra-embryonic membranes that transport oxygen to embryonic tissues. Metabolic rates of embryos from the rigid-shelled eggs of Gehyra variegata did not differ from those of the parchment-shelled eggs of Oedura lesueurii. Moreover, maximum metabolic rates of gekkotans with rigid shells did not differ from those of gekkotan or scincid lizards with parchment shells. In contrast, the yolk sac covered more of the surface area of the egg at oviposition, and the chorioallantois reached its full extent earlier for the species with rigid shelled eggs (Chondrodactylus turneri, G. variegata) than for the species with parchment-shelled eggs (Eublepharis macularius, O. lesueurii). Differences in the temporal patterns of yolk sac and chorioallantois development would thus serve to compensate for low rates of oxygen diffusion through rigid shells of gekkotans.

Water vapor permeability of the rigid-shelled gecko egg.

The vast majority of squamate reptiles (lizards and snakes) produce parchment-shelled eggs that absorb water during incubation, and thus increase in mass, volume, and surface area. In contrast, females from a single monophyletic lineage of gekkotan lizards produce rigid-shelled eggs. These eggs are functionally comparable to those of birds, that is, at oviposition, eggs contain all the water needed for development, and their mass decreases during incubation via the diffusion of water vapor through the shell. I determined patterns of water loss and shell permeability to water vapor from oviposition to hatching for the rigid-shelled eggs of the gekkonid Chrondrodactylus turneri and compared permeability of C. turneri eggs to those of birds and other squamates. Chrondrodactylus turneri eggs incubated at 28.5°C and 40% relative humidity (RH) decreased in mass by 14% over the course of a 68-day incubation period. The rate of water loss varied during incubation; egg mass decreased rapidly during the first 8 days of incubation, declined at a low constant rate during the next 35 days, and then decreased rapidly during the final 25 days of incubation. Overall permeability was 0.17 mg/day/kPa/cm(2) . Percent water loss of rigid-shelled gecko eggs during incubation is similar to that exhibited by birds, but water vapor permeability is about one-third that of bird eggs and several orders of magnitude lower than that of parchment-shelled squamate eggs. In general, the water economy of their eggs may be associated with the adaptive radiation of the rigid-shelled sphaerodactylid, phyllodactylid, and gekkonid geckos.

Hardware implementation algorithm and error analysis of high-speed fluorescence lifetime sensing systems using center-of-mass method.

A new, simple, high-speed, and hardware-only integration-based fluorescence-lifetime-sensing algorithm using a center-of-mass method (CMM) is proposed to implement lifetime calculations, and its signal-to-noise-ratio based on statistics theory is also deduced. Compared to the commonly used iterative least-squares method or the maximum-likelihood-estimation-based, general purpose fluorescence lifetime imaging microscopy (FLIM) analysis software, the proposed hardware lifetime calculation algorithm with CMM offers direct calculation of fluorescence lifetime based on the collected photon counts and timing information provided by in-pixel circuitry and therefore delivers faster analysis for real-time applications, such as clinical diagnosis. A real-time hardware implementation of this CMM FLIM algorithm suitable for a single-photon avalanche diode array in CMOS imaging technology is now proposed for implementation on field-programmable gate array. The performance of the proposed methods has been tested on Fluorescein, Coumarin 6, and 1,8-anilinonaphthalenesulfonate in water/methanol mixture.

Eggs under pressure: components of water potential of chameleon eggs during incubation.

Water exchange of squamate eggs is driven by the difference between the water potentials of eggs and of their nest environment. While osmotic potential is generally assumed to dominate the net water potential of eggs, resistance of the eggshell to stretching also affects egg water potential. We therefore determined osmotic potentials and pressure potentials (mechanical pressure) of eggs of the veiled chameleon Chamaeleo calyptratus over the course of incubation. Because embryos are diapausing gastrulae when eggs are laid and diapause persists several months, the water potential of eggs can be evaluated before it is influenced by the developing embryo. Water uptake during the first 2 wk of incubation was rapid as a result of the large difference between the total water potential of the egg (-848 kPa) and that of its incubation substrate. After about 2 wk, water potential of the egg stabilized at -460 kPa. By day 80 of incubation, the developing embryo and allantois affected water exchange of the egg. The allantoic fluid was initially very dilute, but its osmotic potential decreased to about -200 kPa by the end of incubation. Pressure potential of the egg averaged 25 kPa, with no systematic trend during incubation. The pressure potential exerted by the eggshell reduced the difference between the water potential of the egg and the water potential of the environment, that is, the ability of eggs to take up water. At the time of oviposition, this effect was relatively small, producing a 4%-6% reduction in water potential difference. Once the yolk osmotic potential stabilized, however, the reduction was 12% or more. This observation means that the dynamics of water uptake by squamate eggs cannot be fully understood without consideration of the pressure that is exerted on the contents of eggs by their shells.

Effects of incubation temperature on growth and performance of the veiled chameleon (Chamaeleo calyptratus).

I evaluated the effect of incubation temperature on phenotypes of the veiled chameleon, Chamaeleo calyptratus. I chose this species for study because its large clutch size (30-40 eggs or more) allows replication within clutches both within and among experimental treatments. The major research objectives were (1) to assess the effect of constant low, moderate, and high temperatures on embryonic development, (2) to determine whether the best incubation temperature for embryonic development also produced the "best" hatchlings, and (3) to determine how a change in incubation temperature during mid-development would affect phenotype. To meet these objectives, I established five experimental temperature regimes and determined egg survival and incubation length and measured body size and shape, selected body temperatures, and locomotory performance of lizards at regular intervals from hatching to 90 d, or just before sexual maturity. Incubation temperature affected the length of incubation, egg survival, and body mass, but did not affect sprint speed or selected body temperature although selected body temperature affected growth in mass independently of treatment and clutch. Incubation at moderate temperatures provided the best conditions for both embryonic and post-hatching development. The highest incubation temperatures were disruptive to development; eggs had high mortality, developmental rate was low, and hatchlings grew slowly. Changes in temperature during incubation increased the among-clutch variance in incubation length relative to that of constant temperature treatments.

Developmental arrest during embryonic development of the common chameleon (Chamaeleo chamaeleon) in Spain.

Embryonic development of the common chameleon, Chamaeleo chamaeleon, was monitored from oviposition to hatching at a field site in southwestern Spain and in the laboratory under five experimental temperature regimes. Embryos were diapausing gastrulae at the time of oviposition; developmental arrest in the field continued as cold torpor during winter. Postarrest development in the field commenced in April, and hatching occurred in August, for a total incubation period of 10.5 mo. In the laboratory, one group of eggs was incubated at a constant warm (26 degrees C) temperature. The remaining treatments simulated field conditions and consisted of initial periods of warm temperature of 0, 27, 46, and 71 d, a subsequent 4-mo period of cold winter (16 degrees C) temperature, and a final period of warm (26 degrees C) temperature. Embryos in the constant warm temperature treatment were in diapause an average of 3 mo, with clutch means ranging from 2 to 4 mo. Hatching among clutches occurred over 2 mo. In contrast, for field and experimental eggs that experienced cold winter conditions, hatching within treatments occurred over 2-14 d; "winter" conditions synchronized development. The length of time between the end of cold conditions and hatching did not differ among treatments; development thus resumed as soon as temperature was suitable regardless of the initial period of warm temperature. Diapause in nature thus insures that embryos remain gastrulae after oviposition despite nest temperatures that may be warm enough to support development.

Effects of temperature on embryonic development of the veiled chameleon, Chamaeleo calyptratus.

Temperature dependence of development of the chameleon, Chamaeleo calyptratus, was assessed from observations on eggs incubated at 25, 28 and 30 degrees C. Overall, differentiation, growth in mass, and growth of the yolk sac and chorioallantois were the slowest at 25 degrees C but did not differ between 28 and 30 degrees C. The relative area of the yolk sac (YS), chorioallantoic membrane (CAM), and their precursor, the area opaca vasculosa (AV) was used to characterize developmental phases. During Phase 1, only the AV was present; development was characterized by differentiation with little increase in the size of the embryo. During Phase 2, the vascularized YS and CAM grew from about 10 to 100% coverage of the surface of the shell during a period of about two weeks. Differentiation and growth of the embryo were accordingly rapid. During Phase 3, the YS and CAM were fixed in size and the remainder of development was relatively slow. Characterization of embryonic development with respect to the relative area of the AV-YS-CAM highlighted the functional linkage between development and the systems that provide nutrients to embryos.