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Brooding behavior of the mountain dusky salamander: can the female's presence reduce clutch desiccation?

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The Desmognathus ochrophaeus female decreases clutch desiccation by reducing the exposed surface of her egg mass. -from Author

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... Semiaquatic plethodontids, such as Desmognathus brimleyorum (Trauth 1988), D. fuscus (Hom 1987; Juterbock 1987), and D. ocoee (Forester 1984), attend their eggs by way of body surface contact in nesting conditions typically described as being moisture-laden with a saturated substrate within an enclosed nesting cavity (Forester 1984). Variation, however, exists in the selection of nesting sites among Desmognathus species and presumably reflects some degree of adaptability in desmognathine salamanders to local environmental conditions (Martof and Rose 1963). ...
... Semiaquatic plethodontids, such as Desmognathus brimleyorum (Trauth 1988), D. fuscus (Hom 1987; Juterbock 1987), and D. ocoee (Forester 1984), attend their eggs by way of body surface contact in nesting conditions typically described as being moisture-laden with a saturated substrate within an enclosed nesting cavity (Forester 1984). Variation, however, exists in the selection of nesting sites among Desmognathus species and presumably reflects some degree of adaptability in desmognathine salamanders to local environmental conditions (Martof and Rose 1963). ...
... Our results indicate that female P. albagula oviposit egg clutches in relatively open, exposed nesting perches in the mine shaft; this type of nesting microhabitat is most similar to the cave microenvironment as previously described for this species (Barnett 1970; Noble and Marshall 1929). The assumption that brooding plethodontids possibly reduce the desiccation of their egg clutches during incubation by preventing water loss through body contact with their eggs, as reported for D. ocoee (Forester 1984), appears less plausible for P. albagula. Potential full body contact or near contact through body coiling (our EEBD) around the egg clutch in P. albagula occurred only about 21% of the time. ...
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We examined brooding postures and nest site fidelity in a nesting aggregation of western slimy salamanders (Plethodon albagula) from an abandoned mine shaft located in the Ouachita National Forest of southwestern Arkansas. From November 1999-December 2001, we collected a photographic record of brooding and nesting behavior. Females oviposit a free-hanging, grape-like egg cluster within relatively dry nest perches along the walls of the mine shaft. We recorded six female brooding postures from 101 obser- vations involving 101 egg clutches. The most common brooding posture (34.6% of the time) was one in which the female positioned her shoulder region next to or in contact with her egg clutch. Body coiling around the egg clutch occurred 20.8% of the time and at about the same frequency as brooding postures involving eggs touching the head (17.8%) or trunk (22.8%). Six females (6.3%) exhibited nest site fidelity; one female returned to the same nesting site in each of three nesting seasons, but successfully brooded a clutch during the first season only. Egg predation by a ringneck snake was observed during the 2001 nest- ing season. Our results suggest that brooding postures may function as a predator defense mechanism and may also serve as an anti-microbial defense. The autumn/winter nesting season appears to serve as an anti- predator strategy in this species.
... The eggs of amphibians are protected only by a semi-permeable jelly coat and are therefore vulnerable to continuous threats of desiccation, fungal infections, and predation throughout embryonic development (Salthe and Mecham 1974). Previous studies have shown that brooding behaviors increase egg survival via several mechanisms: reducing predation (Highton and Savage 1961;Forester 1983); increasing hydration (Forester 1984;Taigen et al. 1984); and reducing fungal or bacterial infection (Forester 1981;Austin 2000). We hypothesize that brooding behavior of larger females can also affect specific offspring traits such as offspring size. ...
... In addition, cross-foster and control females did not significantly differ in the rate of abandonment or in the number of eggs eaten throughout development. Abandonment and oophagy are common in field populations of P. cinereus (Crespi 2001; J. Gillette, K. Yurewicz, and H. Wilbur, personal communication) and have been documented in other lab experiments with brooding plethodontid salamanders (Highton and Savage 1961;Forester 1984). Even with reduced sample sizes, and therefore reduced statistical power, our ability to detect significant correlations in this study suggests that the effect of the parenting mother's body size is robust. ...
... Unfortunately, the exact mechanism through which these behaviors enhance development could not be determined by this experiment. The positive effect of contact time with eggs may be due to the reduction of desiccation or suppression of infections from microorganisms (see Forester 1984;Austin 2000). It is interesting that the number of times mothers moved eggs was negatively correlated to offspring size since vibration of eggs (simulation of gular movements of the mother) has been shown to enhance egg survival (Forester 1984). ...
Article
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In the terrestrial salamander (Plethodon cinereus), previous work has shown that mother's body size is positively correlated to offspring size at the time of hatching even after controlling for the effects of egg size. This study was designed to determine whether maternal body size affects offspring size via pre-oviposition factors (e.g., yolk quality, jelly coat composition, or maternal genes) or post-oviposition factors (e.g., parental care behaviors, parental modification of environment). Gravid females were captured and induced to lay eggs in experimental chambers in which the environment was standardized. Fifteen clutches were exchanged, or cross-fostered, between female pairs differing in body size. Ten females whose eggs were taken away and then returned served as controls for the crossing treatment. Foster mothers did not significantly differ from control mothers in the time spent with eggs, body position, or number of egg movements during brooding. Average egg mass measured midway through development was not significantly correlated to the body size of either the genetic or foster mother, but was correlated to pre-oviposition oocyte size. At hatching, offspring body length was positively correlated to egg size and the foster mother's body size. This correlation suggests that in P. cinereus post-oviposition maternal effects have a greater impact on offspring size than other maternal factors incorporated into the egg prior to oviposition. While our study showed that larger mothers moved their eggs less often and tended to spend more time in contact with their eggs, further work needs to be done to identify the specific mechanisms through which larger mothers influence the body size of their offspring. This is the first experimental demonstration of post-oviposition maternal effects for any amphibian with parental care.
... Egg attendance, whereby a parent remains with the egg mass at a fixed location, represents by far the most common mode of amphibian parental care (Wells 1981(Wells , 2007Crump 1996). By investing in egg attendance, amphibians can increase offspring survivorship by reducing or preventing developmental abnormalities (Forester 1979;Simon 1983), predation (Forester 1979;Kluge 1981;Townsend et al. 1984;Junc a 1996;Bickford 2004;Vockenhuber et al. 2009), and desiccation (Weygoldt 1980;Forester 1984;Taigen et al. 1984;Bickford 2004;Delia et al. 2013). In addition, it has been suggested that adults may reduce fungal infection of embryos during attendance (Salthe & Mecham 1974;Simon 1983;Green 1999). ...
... This is suggested by the results in adult removal experiments showing high levels of egg desiccation in the absence of attending adults, whereas control clutches were buffered from this source of mortality (Fig. 3), supporting Wilson's (1975) hypothesis that harsh environments can be a driver of parental care. Prevention of water loss has been cited as a major function of egg attendance in amphibians, and it has been suggested that adults may (1) reduce surface exposure and thus evaporation in embryos (Forester 1984), (2) move water osmotically from their ventral integument to embryos (Taigen et al. 1984;Bickford 2004), (3) bring water to embryos by repeatedly soaking themselves in water and placing their bodies over clutches (Ueda 1986;Cheng & Kam 2010), and (4) actively release internal fluids over embryos to increase hydration (Weygoldt 1980(Weygoldt , 1987Bourne 1998). Observations of C. hansenae indicate that it may be using a combination of these methods, including reducing surface exposure by physically covering the clutch and possibly secreting liquids to maintain or increase hydration of embryos. ...
Article
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The arboreal frog, Chiromantis hansenae (Family: Rhacophoridae), is one of only a handful of South‐East Asian amphibian species reported with parental care. We present the first systematic observational and experimental study confirming offspring benefits as a result of this care, which has a number of unusual life‐history characteristics. Eggs are unusually small, breeding takes place in large pools, and females attend the eggs. Field observations and an adult removal experiment demonstrated a critical contribution of egg attendance to offspring survivorship. Harsh environmental conditions for offspring appeared to be the prime mover of parental care in this species, with desiccation as the main source of mortality when attending adults are absent. Results confirm females to be the caregivers, making C. hansenae a rare case of maternal egg attendance in a non‐directly developing anuran.
... As in many salamanders, D. ochrophaeus brood their clutches until hatching, and shortly thereafter . Clever experiments in a few species have shown that mothers improve hatching success via several mechanisms (reviews: Forester, 1979; Nussbaum, 1985; Horn et ai, 1990 ). First, mother's presence both reduces desiccation (Forester, 1984 ) and deters most clutch predation (Forester, 1979Forester, , 1983 ). Second, females physically manipulate clutches which is thought both to promote gas exchange and to inhibit growth of fungal mycelia because unattended clutches succumb to fungal infection and death in a few days (Forester, 1979; Bernardo, personal observation). ...
... Despite the fitness gains that mothers realize by brooding, they are also subject to personal costs. Attending females do not actively feed, and they lose mass both to dehydration and tissue loss during brooding (Forester, 1981Forester, , 1984). Brooding females are susceptible to predation by larger vertebrate predators (Forester, 1979 ). ...
Article
SYNOPSIS. Maternal effects comprise a class of phenotypic effects that parents have on phenotypes of their offspring that are unrelated to the offspring's own genotype. Although biologists have known about the importance of these effects for decades, maternal effects have only recently been studied widely by evolutionary ecologists. Moreover, the impact of maternal effects on the rate and direction of evolution of other components of the phenotype has only recently been fully elucidated by theoretical genetic models. This paper provides a brief overview of maternal effects research, focusing on research conducted in animal systems. First, I review and summarize definitions and theoretical treatments of maternal effects. Then I survey the diversity of maternal effects and some of the kinds of ecological and evolutionary impacts that maternal effects have been shown to have on offspring performance. Third, I discuss some of the ecological and evolutionary implications of maternal effects in ecological research. In this context, it is argued that the kinds of maternal effects that have been studied, and some of the potential implications of maternal effects that have not been explored are both artifacts of research effort. Hence, I identify several fruitful areas for future maternal effects research. Lastly, I describe the necessity of explicitly incorporating maternal effects in ecological research, whether or not the researcher is focusing on them as the topic of the research.
... This form of egg attendance, or brooding behavior, consists of the parent covering the eggs with their body to reduce water loss and/or directly releasing liquid to the eggs through the venter or from their cloaca (Crump 1996). In some lungless salamanders (Plethodontidae), unattended eggs lose more water per hour than eggs attended by a female (Forester 1984). Also, in glass frogs (Centrolenidae), egg attendance positively affects hydration, and parents adjust the frequency of attendance to both weather conditions and egg dehydration status (Delia et al. 2013). ...
Article
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Parenting is considered a key evolutionary innovation that contributed to the diversification and expansion of vertebrates. However, we know little about how such diversity evolved. Amphibians are an ideal group in which to identify the ecological factors that have facilitated or constrained the evolution of different forms of parental care. Among, but also within, the three amphibian orders—Anura, Caudata, and Gymnophiona—there is a high level of variation in habitat use, fertilization mode, mating systems, and parental sex roles. Recent work using broad phylogenetic, experimental, and physiological approaches has helped to uncover the factors that have selected for the evolution of care and transitions between different forms of parenting. Here, we highlight the exceptional diversity of amphibian parental care, emphasize the unique opportunities this group offers for addressing key questions about the evolution of parenting, and give insights into promising novel directions of research. Expected final online publication date for the Annual Review of Ecology, Evolution, and Systematics, Volume 54 is November 2023. Please see http://www.annualreviews.org/page/journal/pubdates for revised estimates.
... The adaptive benefits of plethodontid maternal behaviors are diverse (Forester 1979). Brooding females provide active and passive defense against conspecific and heterospecific predators (Highton and Savage 1961;Bachmann 1964; but see Carreño and Harris 1998), move and turn their eggs to increase oxygenation and prevent adhesive malformations (Stebbins 1954;Vandel and Bouillon 1959;Durand and Vandel 1968;Salthe and Mecham 1974), and moisten eggs during periods of drought (Forester 1984). Furthermore, mothers ingest infected eggs to prevent the spread of fungal infection (Snyder 1971;Tilley 1972;Salthe and Mecham 1974). ...
... The adaptive benefits of plethodontid maternal behaviors are diverse (Forester 1979). Brooding females provide active and passive defense against conspecific and heterospecific predators (Highton and Savage 1961;Bachmann 1964; but see Carreño and Harris 1998), move and turn their eggs to increase oxygenation and prevent adhesive malformations (Stebbins 1954;Vandel and Bouillon 1959;Durand and Vandel 1968;Salthe and Mecham 1974), and moisten eggs during periods of drought (Forester 1984). Furthermore, mothers ingest infected eggs to prevent the spread of fungal infection (Snyder 1971;Tilley 1972;Salthe and Mecham 1974). ...
Article
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Despite rising interest among scientists for over two centuries, parental care behavior has not been as thoroughly studied in amphibians as it has in other taxa. The first reports of amphibian parental care date from the early 18th century, when Maria Sibylla Merian went on a field expedition in Suriname and reported frog metamorphs emerging from their mother's dorsal skin. Reports of this and other parental behaviors in amphibians remained descriptive for decades, often as side notes during expeditions with another purpose. However, since the 1980s, experimental approaches have proliferated, providing detailed knowledge about the adaptive value of observed behaviors. Today, we recognize more than 30 types of parental care in amphibians, but most studies focus on just a few families and have favored anurans over urodeles and caecilians. Here, we provide a synthesis of the last three centuries of parental care research in the three orders comprising the amphibians. We draw attention to the progress from the very first descriptions to the most recent experimental studies, and highlight the importance of natural history observations as a source of new hypotheses and necessary context to interpret experimental findings. We encourage amphibian parental care researchers to diversify their study systems to allow for a more comprehensive perspective of the behaviors that amphibians exhibit. Finally, we uncover knowledge gaps and suggest new avenues of research using a variety of disciplines and approaches that will allow us to better understand the function and evolution of parental care behaviors in this diverse group of animals.
... Their contributions to synthesizing our understanding of amphibian and reptile behaviors cannot be understated. Thanks to P. Zani for reviewing Egg attendance in these species is known to reduce desiccation and predation risk, [174][175][176][177] and females will often consume dead or infected eggs to prevent the spread of fungi to other embryos. 175 It is also hypothesized that female contact with eggs may facilitate the transfer of beneficial microbes that inhibit fungal colonization. ...
... Female terrestrial plethodontid salamanders oviposit clutches in moist, protected terrestrial sites where they brood the eggs for several months (Nussbaum 1985). Egg brooding increases egg survival by reducing predation, yolk layering, fungal infection, and desiccation (Forester 1984, Highton and Savage 1961, Synder 1971. Egg guarding, however, can be a costly investment for females. ...
Preprint
In some species of vertebrates egg brooding is a costly form of parental care. Therefore, misdirection of parental care can significantly lower a female's fitness. Because of the maternal investment and increased survivorship to offspring from egg guarding, a brooding female should home to her nest site after being displaced a short distance and discriminate between her own eggs and eggs from other females. In this study, we experimentally tested, in the field, alternative hypotheses concerning homing ability and egg discrimination in a population of nesting western slimy salamanders ( Plethodon albagula ). Fourteen brooding females were displaced 1 m to the left or right of their nest sites (determined randomly) for the homing experiment. Furthermore, brooding females (n = 13) were presented with their own clutches, which were displaced 50 cm to the left or right (determined randomly), and unfamiliar egg clutches at their original nest sites. The females were released at an equal distance from both egg clutches. After 24 hours, 12 displaced females (86%) had returned to their own nest sites and were brooding their egg clutches. Also, after 24 hours, nine test females had returned to their own nest sites and were brooding the unfamiliar egg clutches. No control or test females were present at the other new nest site locations. Therefore, we suggest that brooding female P. albagula do home to their nest sites and exhibit indirect egg discrimination.
... Chen et al. (2007) reported that attending male K. eiffingeri spent 1/3 of their time brooding eggs. Brooding behavior was demonstrated to serve the purpose of preventing egg desiccation, either through osmosis or through decreasing the exposed surface area of a clutch (Wells 1981, Forester 1984, Taigen et al. 1984, Townsend et al. 1984, Bickford 2004. Egg attendance by male frogs is likely to reduce the chance of eggs being eaten by small insects such as ants, earwigs, and aphids (Kam et al. 1996, Chen et al. 2007. ...
Article
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We conducted a field study in Mar.-June 2007 to evaluate the importance of egg attendance by male Kurixalus eiffingeri to egg survivorship. We found 38 egg clutches and visited them on a fixed scheme by checking all nests at night for 6 d, yielding 1368 visits of 10 min each. Egg attendance frequency was highly uneven among nests and was positively correlated with clutch size. On average, 6.8% ± 4.4% of eggs in an egg clutch died each day. Daily egg mortality was inversely correlated with the attendance frequency, suggesting that paternal care is important to egg survival even though this is a low nest-attendance species. Variances of egg mortality were low when paternal care effort was high, but opposite results were found when paternal care effort was low, suggesting the fate of eggs can also be affected by factors other than paternal care, such as stump conditions. Our results suggest that parental care effort varies via a cost-benefit relationship of investment in the present offspring vs. future reproductive chances. Male frogs spent more time attending eggs when the clutch sizes were large but sought additional mating opportunities if the egg clutches were small, which resulted in large variations in egg mortality among clutches.
... Females of most terrestrial plethodontids brood their clutches for several months, during which time they do not feed (Organ 1961, Forester 1984. Brooding probably prevents desiccation and fungal infections of eggs, and provides protection against predators, including other salamanders (Highton and Savage 1961, Forester 1979, Bachman 1984. ...
Article
ABSTRACT The Clouded Salamander, Aneides ferreus, is a terrestrial forest-dwelling salamander with a distribution in British Columbia limited to Vancouver Island and some surrounding small islands. It is typically found under the bark or within cracks of decomposing logs, under loose bark and other debris on the ground, and in talus. Courtship and mating take place in the spring and females lay small clutches of terrestrial eggs in cavities in decomposing logs in the early summer. The larval stage is passed in the egg,
... Brooding females stop feeding and seldom venture from nest sites, except in response to large predators or seasonal flooding (Forester 1981(Forester , 1983Juterbock 1987;Hom 1987). Brooding behaviour contributes to egg survival by reducing egg susceptibility to predation (Bachmann 1964;Snyder 1971;Forester 1978Forester , 1979a, yolk layering (Forester 1979a), fungal infestation (Snyder 1971;Forester 1979 a), and desiccation (Forester 1984), but the cost to the female is high. Egg production and brooding have been estimated to represent 48 % of a female's annual energy budget (Fitzpatrick 1973). ...
Article
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Egg guarding, or `brooding', by the mountain dusky salamander (Desmognathus ochrophaeus) is an example of maternal behaviour that can be experimentally manipulated and described quantitatively. It has been demonstrated that females of this species can specifically recognize, and will preferentially brood, their own eggs over those of a conspecific. We investigated whether this behaviour would extend to the selection of eggs of a more genetically similar animal in preference to those of a less similar animal, as might be predicted by inclusive fitness theory. We report here a highly significant correlation between time spent brooding and genetic similarity determined by random amplification of polymorphic DNA (RAPD) analysis. This is the first demonstration of a quantitative relation between genetic relatedness and maternal care in amphibians. Our findings have implications for the nature of maternal kin recognition in amphibians and its effect on kin selective behaviour in this class.
... This is surprising because D. ocoee is one of the most common salamanders in the region (Smith & Petranka, 2000), and it has figured prominently as a model system for many other studies (Bernardo, 2003). Although a great deal is known about oviposition behaviour, clutch size and brooding behaviour in D. ocoee (Tilley, 1972;Forester, 1979aForester, ,b, 1981Forester, , 1983Forester, , 1984Forester, , 1986, and much has been learned about its size at metamorphosis (Bernardo & Reagan-Wallin, 2002), juvenile growth and development during the terrestrial stage, survival and maturation (Tilley, 1977(Tilley, , 1980Bernardo, 1994 and unpublished), its larval life history has been largely unstudied. A brief summary of the generalized life cycle of D. ocoee (Fig. 2) aids in understanding the phenology of the larval stage and thus, the logic and goals of the sampling scheme used in the present study. ...
Article
Recent models argue that thermal environments are the major cause of ectotherm life-history clines. However, elevational clines in body size in the mountain dusky salamander Desmognathus ocoee (family Plethodontidae) shift from positive at hatching, to negative at metamorphosis to positive again as adults, and so are not consistent with this explanation. The clinal shift from hatching to metamorphosis was investigated by examining the clinal and seasonal feeding patterns of larval salamanders at high and low elevation sites in rockface and woodland habitats. Repeated cohort sampling was also used to examine clinal and seasonal patterns in body size and to estimate average growth rates. Larval growth in both rockface and woodland habitats was tightly correlated with feeding activity. Although temperature was found to vary between high and low elevation sites, the greatest growth occurred in a cold woodland habitat with a high elevation, and the lowest growth occurred in an adjacent rockface habitat. Because this difference in growth cannot be attributed to thermal differences, we conclude that local food resource levels are the predominant source of local differences in growth. These findings, clinal patterns of variation in other predatory salamanders, and experimental analyses in which both food and temperature are orthogonally manipulated, indicate that general models that single out temperature as the principle cause of ectotherm life-history clines should be viewed with caution.
... Often associated with male provision of a nest site is male parental (paternal) care, during which time the male may stay with the eggs during part, or all, of the eggs' development (Crump 1995). While in attendance, adult males can protect the brood against predators (Simon 1983), desiccation (Forester 1984), fungal infestation (Forester 1979), and developmental abnormalities (Simon 1983). ...
Article
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We examined multiple mate choice criteria in Cophixalus ornatus, a terrestrial breeding, microhylid frog. Mate choice consisted of three stages: mate attraction (male calling), courtship (male behavior between the call site and the nest), and nest site selection by the female. For male C. ornatus, the possession of a call with low dominant frequency relative to calling neighbors increased the probability that they would attract females. Dominant frequency was negatively correlated with age independent of male mass and snout vent length. When escorting the female from the call site to their nest, males traveled along more convoluted paths than when returning to the nest alone. The convolution of the path was, therefore, considered an aspect of courtship. Females released eggs into nests with structural characteristics typical of nests constructed by older males. Thus, females increased their chances of locating an acceptable nest by preferentially approaching males with lower dominant frequencies. This study is the first to demonstrate that age, independent of mass or snout-vent length, can influence call characteristics in anurans, and it is also the first to demonstrate the importance of male age to female mate choice in an amphibian.
... In E. coqui, water-conserving postures prevented potentially lethal rates of dehydration, which allowed frogs to forage in highly variable conditions of the forest canopy ( Pough et al., 1983). Furthermore, demognathine salamanders are reported to decrease clutch desiccation by coiling tightly about their eggs (Forester, 1984). ...
Article
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Bothrops moojeni is a member of the atrox group that occurs in central and southeastern Brazil and adjacent Paraguay and Argentina. We describe habitat use, diel and seasonal activity, biometry, feeding habits, and reproduction of B. moojeni, based on field studies and analysis of 207 preserved specimens. Bothrops moojeni predominantly uses riparian vegetation in the Cerrado (central Brazilian savannas), such as gallery forests and adjacent wet grasslands, although they occasionally use drier interfluvial areas. Sexes are dimorphic in snout-vent length, relative tail length, relative mass, and relative head length. Main prey types were mammals, frogs, and lizards. There is an ontogenetic dietary shift from ectotherms to endotherms. Prey- predator mass ratio ranged from 0.008-1.079, and prey mass was positively correlated with predator mass. Although prey absolute mass was positively correlated with snake mass, large snakes did not eliminate small prey from their diets. There was an ontogenetic decrease in prey relative mass. Females consumed more endothermic prey than males, and this difference probably reflects size differences. The reproductive cycle is lengthy and seasonal, with recruitment restricted to the rainy season. Fecundity is positively correlated with female body size. Life-history characters of B. moojeni are similar to those of other members of the atrox group, even though B. moojeni occurs in the Cerrado, a drier and more seasonal biome. Bothrops moojeni is a large pitviper from riparian areas in central and southeastern Brazil, throughout the Cerrado morphoclimatic domain (Borges and Araujo, 1998; Campbell and Lamar, 1989; Leloup, 1984). Recent systematic studies indicate that populations of B. moojeni from the Cerrado of southeastern and central Brazil form a monophyletic assemblage within the atrox spe-
... We observed in the field that male C. eiffingeri spent more than 1/3 of their time actively moistening clutches with their ventral surface, suggesting that brooding is the major component of egg attendance. In many terrestrial and arboreal frog species, brooding behavior has been demonstrated to prevent egg desiccation, either through osmosis or through decreasing the exposed surface area of the clutch (Wells, 1981;Forester, 1984;Taigen et al., 1984;Townsend et al., 1984;Bickford, 2004). ...
Article
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Field observations were conducted on egg attendance in Chirixalus eiffingeri from April to August 2003. Parental attendance during embryonic development was performed exclusively by males. The frequency of egg attendance was low (27%), but it had a distinct diel pattern in which males were observed to attend eggs more frequently at night than during the day. Attendance frequency significantly decreased with increasing developmental stage of the embryos, but it was not statistically significantly related to clutch size. Field observations confirmed that male frogs actively moisten egg clutches using their ventral surfaces, presumably to prevent desiccation of egg clutches. The non-significant relationship between hatching success and frequency of egg attendance suggests that embryonic survival of C. eiffingeri is more than a function of egg attendance, and ecological and environmental factors, such as climate and characteristics of microhabitats, may also influence the survivorship of the embryos.
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A review of systematics, taxonomy, genetics, biogeography and natural history of the genus Speleomantes Dubois, 1984 (Amphibia Caudata Plethodontidae). Outside the American continent the family Plethodontidae includes only the south Korean Karsenia koreana Min,Yang, Bonett, Vieltes, Brandon et Wake, 2005, and the European genus Speleomantes Dubois, 1984, whose systematics and taxonomy have rather recently been object of a morphologic and genetic review, which allowed to recognize 3 continental species [S. strinatii (Aellen, 1958); S. ambrosii ambrosii (Lanza, 1955) and S. ambrosii bianchii Lanza, Cimmaruta, Forti, Bullini et Nascetti, 2005; S. italicus (Dunn, 1923)] and 4 Sardinian ones [ S. flavus (Stefani, 1969); S. supramontis (Lanza, Nascetti et Bullini, 1986); S. imperialis imperialis (Stefani, 1969) and S. imperialissarrabusensis Lanza, Leo, Forti, Cimmaruta, Caputo et Nascetti, 2001); S. genei (Temminck et Schlegel, 1838) with subsp A and subsp. B]. Until recently the knowledge of Speleomantes’ ecoethology has been widely lacunose and specific research on this topic started only in the nineties. The present paper summarizes the current knowledge onthe European plethodontid salamanders, genus Speleomantes. The authors deal with systematics, taxonomy, geographical and altitudinal distribution, genetics (two keys to species and subspecies are given: one based on morphology and geographical distribution, the other based on genetic characters), biogeography, ecology (habitats, herpetocoenosis, predators, parasites, food), ethology (feeding behaviour, activity, habitat use, displacement, antipredator adaptations, communication), reproduction (sexual dimorphism, gametogenesis, mating behaviour, oviparity, ovovi-viparity, parental cares), development, population dynamics, conservation (abundance, threats, conservation guidelines), as well as with research prospects. The quoted bibliography includes 565 titles. Key words: Amphibia, Plethodontidae, Speleomantes, systematics, taxonomy, keys, genetics, biogeography, biology,conservation, France, Italy. Revisione della sistematica, tassonomia, genetica, biogeografia e storia naturale del genere Speleomantes Dubois, 1984 (Amphibia Caudata Plethodontidae). Al di fuori del continente americano la famiglia Plethodontidae conta solo la specie sud coreana Karsenia koreana Min, Yang, Bonett, Vieltes, Brandon et Wake, 2005, e il genere europeo Speleomantes Dubois, 1984, la cui sistematica e tassonomia, oggetto di una relativamente recente revisione su basi genetiche e morfologiche, ha portato a riconoscere 3 specie continentali [S. strinatii (Aellen, 1958); S. ambrosii ambrosii (Lanza, 1955) e S. ambrosii bianchii Lanza, Cimmaruta, Forti, Bullini et Nascetti, 2005; S. italicus (Dunn, 1923)] e 4 sarde [S. flavus (Stefani, 1969); S. supramontis (Lanza, Nascetti et Bullini, 1986); S. imperialis imperialis (Stefani, 1969) e S. imperialis sarrabusensis Lanza, Leo, Forti, Cimmaruta, Caputo et Nascetti, 2001); S. genei (Temminck et Schlegel, 1838) con le sottospecie A e B]. Sino a poco tempo fa le conoscenze sull’ecoetologia dello Speleomantes sono rimaste ampiamente lacunose e ricerche approfondite in proposito sono state avviate solo a partire dagli anni ‘90. Il presente contributo riassume le attuali conoscenze sui Pletodontidi europei del genere Speleomantes. Gli autori trattano sistematica, tassonomia, distribuzione geografica ed altitudinale, genetica (vengono fornite due chiavi per l’identificazione delle specie e sottospecie: una basata sulla morfologia e la distribuzione geografica, l’altra basata sulle caratteristiche genetiche), biogeografia, ecologia (habitat, erpetocenosi, predatori, parassiti, dieta), etologia (comportamento predatorio, attività, uso dell’habitat, spostamenti, adattamenti antipredatori, comunicazione), riproduzione (dimorfismo sessuale, gametogenesi, corteggiamento, oviparità, ovoviviparità e cure parentali), sviluppo, dinamica di popolazione, conservazione (abbondanza, minacce, linee guida per la conservazione) e prospettive di ricerca. La bibliografia citata comprende 565 titoli. Parole chiave: Amphibia, Plethodontidae, Speleomantes, sistematica, tassonomia, chiavi, genetica, biogeografia, biologia, conservazione, Francia, Italia. (10) (PDF) A review of systematics, taxonomy, genetics, biogeography and natural history of the genus Speleomantes Dubois, 1984 (Amphibia Caudata Plethodontidae). Available from: https://www.researchgate.net/publication/292730134_A_review_of_systematics_taxonomy_genetics_biogeography_and_natural_history_of_the_genus_Speleomantes_Dubois_1984_Amphibia_Caudata_Plethodontidae [accessed Mar 09 2023].
Article
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Chapter
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Chapter
Egg brooding behavior is typical in plethodontid salamanders, and in most species females attend their eggs from oviposition until hatching (Crump, 1995). Attendant behavior has been shown to increase egg survival in the field (Forester, 1979a;Harris et al., 1995), and it has been specifically demonstrated to reduce susceptibility of eggs to predation (Bachmann, 1984;Forester, 1979a,Forester, 1979a), yolk layering (Forester, 1979a), fungal infestation (Austin, 1997;Forester, 1979a;Ng and Wilbur, 1995;Snyder, 1971), and desiccation (Forester, 1984;Ng and Wilbur, 1995). Such behavior, however, represents a significant investment for the female, resulting in weight loss (Forester, 1981;Harris et al., 1995;Ng and Wilbur, 1995) and increased exposure to predators (Hom, 1988).
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