Multiple Paternity Benefits Female Marbled Salamanders by Increasing Survival of Progeny to Metamorphosis

Multiple paternity occurs in most species and animal groups that have been studied. Because mating involves fitness costs to individual females, theory predicts that polyandrous females gain greater fitness benefits than costs, allowing the behavior to be maintained. Genetic, rather than material, benefits often occur in species where males provide females with little more than sperm and seminal fluid. We compared fitness correlates of single‐ and double‐sire clutches from female marbled salamanders (Ambystoma opacum) at the egg, hatchling, and metamorph stages of offspring development. Because clutches were collected from experimental breeding groups, strict paternity exclusion of offspring using microsatellite data allowed us to categorize each clutch as having either one or two fathers. Early offspring viability and size of hatchlings were not different between single‐ and multiple‐paternity clutches. Larvae from the two clutch types were allowed to develop together in field enclosures until metamorphosis. Although there was no difference in size at metamorphosis, survival to metamorphosis was significantly higher in multiple‐paternity clutches (44% vs. 40%) suggesting a benefit for females. The results were consistent with genetic benefits, although maternal effects could not be ruled out. The data did not support predictions of the genetic bet‐hedging and good sperm hypotheses for genetic benefits of polyandry.     

Male Courtship Pheromones Induce Cloacal Gaping in Female Newts (Salamandridae)

Pheromones are an important component of sexual communication in courting salamanders, but the number of species in which their use has been demonstrated with behavioral evidence remains limited. Here we developed a behavioral assay for demonstrating courtship pheromone use in the aquatically courting Iberian ribbed newt Pleurodeles waltl. By performing an in-depth study of the courtship behavior, we show that females invariably open their cloaca (cloacal gaping) before engaging in pinwheel behavior, the circling movement that is the prelude to spermatophore uptake. In contrast, cloacal gaping was not observed in failed courtships, where females escaped or displayed thanatosis. Since gaping mainly occurred during male amplexus and cloacal imposition, which is the obvious period of pheromone transfer, we next investigated whether male courtship water (i.e., water holding courtship pheromones) alone was able to induce this reaction in females. These tests showed that courtship water induced cloacal gaping significantly more than water, even in the absence of a male. Cloacal gaping thus provides a simple and robust test for demonstrating courtship pheromone use in the Iberian ribbed newt. Since opening the cloaca is an essential prerequisite for spermatophore pick-up in all internally fertilizing salamanders, we hypothesize that variations on this assay will also be useful in several other species.     

Ovarian Hormones Influence Territorial Aggression in Free-Living Female Mountain Spiny Lizards

Females are aggressive in many species but relatively little is known about the hormonal basis of female aggression, especially in free-living animals. Female mountain spiny lizards aggressively defend territories from other females. Previously, we showed that plasma levels of testosterone (T) and estradiol (E) are positively associated with levels of female aggression. Here, we manipulated hormone levels in free-living females and examined aggression expressed by females returned to their natural territories. Females received one of the following: (1) ovariectomy + empty implant (OVEX), (2) ovariectomy + T implant (T-IMP), or (3) sham surgery + empty implant (SHAM). OVEX females had reduced plasma levels of E but not T relative to SHAM females. T-IMP females had elevated plasma levels of T. Levels of display and aggression in OVEX females were reduced relative to SHAM females. T-IMP females had restored levels of display behavior although, unlike SHAM, no T-IMP females expressed the overt aggressive behavior of charging. These data are most consistent with the hypothesis that an ovarian factor such as E promotes female aggression, since ovariectomy reduced both plasma E and aggression but had no effect on plasma T. The results from the T-IMP females are also consistent with this hypothesis if we assume that the effects of T are due to aromatization to E in target tissues. The data do not rule out a role for T in promoting female aggression since T-implants resulted in elevated plasma T and restored display behaviors. This study represents one of the first studies examining the hormonal basis of female aggression in free-living females.     

Male Courtship Pheromones Affect Female Behaviour in the Swordtail Characin (Corynopoma riisei)

Pheromones constitute an important cue used by both males and females during courtship. Here, we investigate the effect of male pheromones on female behaviour in the swordtail characin (Corynopoma riisei), a species of fish where males have a caudal pheromone gland which has been suggested to affect female behaviour during courtship. We subjected female C. riisei to male courtship pheromones and investigated the effect on both female behaviour and brain serotonergic activity levels compared to a control group. While no difference in serotonergic activity was found, the pheromone‐treated females showed lower stress levels compared to the control group. Furthermore, pheromone‐treated females increased locomotor activity over time, while a decrease in locomotor activity was observed in the control group. These results suggest that the male courtship pheromones may serve to reduce female stress and increase female activity, possibly to aid males in gaining access to females and facilitating sperm transfer.     

Evolution of Gigantism in Amphiumid Salamanders

The Amphiumidae contains three species of elongate, permanently aquatic salamanders with four diminutive limbs that append one, two, or three toes. Two of the species, Amphiuma means and A. tridactylum, are among the largest salamanders in the world, reaching lengths of more than one meter, whereas the third species (A. pholeter), extinct amphiumids, and closely related salamander families are relatively small. Amphiuma means and A. tridactylum are widespread species and live in a wide range of lowland aquatic habitats on the Coastal Plain of the southeastern United States, whereas A. pholeter is restricted to very specialized organic muck habitats and is syntopic with A. means. Here we present analyses of sequences of mitochondrial and nuclear loci from across the distribution of the three taxa to assess lineage diversity, relationships, and relative timing of divergence in amphiumid salamanders. In addition we analyze the evolution of gigantism in the clade. Our analyses indicate three lineages that have diverged since the late Miocene, that correspond to the three currently recognized species, but the two gigantic species are not each other''s closest relatives. Given that the most closely related salamander families and fossil amphiumids from the Upper Cretaceous and Paleocene are relatively small, our results suggest at least two extreme changes in body size within the Amphuimidae. Gigantic body size either evolved once as the ancestral condition of modern amphiumas, with a subsequent strong size reduction in A. pholeter, or gigantism independently evolved twice in the modern species, A. means and A. tridactylum. These patterns are concordant with differences in habitat breadth and range size among lineages, and have implications for reproductive isolation and diversification of amphiumid salamanders.     

Acoustic Monitoring Reveals Congruent Patterns of Territorial Singing Behaviour in Male and Female Tropical Wrens

Many animals defend territories against conspecific individuals using acoustic signals. In birds, male vocalizations are known to play a critical role in territory defence. Territorial acoustic signals in females have been poorly studied, perhaps because female song is uncommon in north‐temperate ecosystems. In this study, we compare male vs. female territorial singing behaviour in Neotropical rufous‐and‐white wrens Thryothorus rufalbus, a species where both sexes produce solo songs and often coordinate their songs in vocal duets. We recorded free‐living birds in Costa Rica using an eight‐microphone Acoustic Location System capable of passively triangulating the position of animals based on their vocalizations. We recorded 17 pairs of birds for 2–4 consecutive mornings and calculated the territory of each individual as a 95% fixed kernel estimate around their song posts. We compared territories calculated around male vs. female song posts, including separate analyses of solo vs. duet song posts. These spatial analyses of singing behaviour reveal that males and females use similarly sized territories with more than 60% overlap between breeding partners. Territories calculated based on solo vs. duet song posts were of similar size and similar degrees of overlap. Solos and duets were performed at similar distances from the nest for both sexes. Overall, male and female rufous‐and‐white wrens exhibit very similar spatial territorial singing behaviour, demonstrating congruent patterns of male and female territoriality.     

Love Is Blind: Indiscriminate Female Mating Responses to Male Courtship Pheromones in Newts (Salamandridae)

Internal fertilization without copulation or prolonged physical contact is a rare reproductive mode among vertebrates. In many newts (Salamandridae), the male deposits a spermatophore on the substrate in the water, which the female subsequently takes up with her cloaca. Because such an insemination requires intense coordination of both sexes, male newts have evolved a courtship display, essentially consisting of sending pheromones under water by tail-fanning towards their potential partner. Behavioral experiments until now mostly focused on an attractant function, i.e. showing that olfactory cues are able to bring both sexes together. However, since males start their display only after an initial contact phase, courtship pheromones are expected to have an alternative function. Here we developed a series of intraspecific and interspecific two-female experiments with alpine newt (Ichthyosaura alpestris) and palmate newt (Lissotriton helveticus) females, comparing behavior in male courtship water and control water. We show that male olfactory cues emitted during tail-fanning are pheromones that can induce all typical features of natural female mating behavior. Interestingly, females exposed to male pheromones of their own species show indiscriminate mating responses to conspecific and heterospecific females, indicating that visual cues are subordinate to olfactory cues during courtship.     

Effects of Timber Management on Pond-Breeding Salamanders

Abstract: Pond-breeding salamanders spend most of their lives in forested habitat surrounding the vernal pools where they breed. Timber harvesting has been demonstrated to have negative impacts on salamander populations due to changes in soil temperature, soil compaction, and general degradation of habitat. However, little is known about how long it takes for harvested forest habitat to once again become suitable for salamanders. Questions also remain as to whether salamanders will use an area that has been harvested in recent years if an older intact forest area is available. We used drift fences and pitfall traps to capture adult spotted salamanders (Ambystoma maculatum) and marbled salamanders (A. opacum) migrating to 3 vernal ponds during their breeding seasons. The study area contained tracts of forest that were clear-cut 11–12 years prior to the study. All 3 ponds were surrounded by areas of clear-cut and intact forest and drift fences were placed in both habitat types. Similar numbers of spotted salamanders entered the ponds from clear-cut and intact forest areas. The number of marbled salamanders migrating to the ponds did not differ between areas of clear-cut and intact forest. These results suggest that clear-cut habitats may become suitable for adult pond-breeding salamanders after a relatively short regeneration period.     

Male-male Competition and Female Mate Choice through Courtship Display in the Territorial Damselfish Stegastes nigricans

Both sexes of the herbivorous damselfish Stegastes nigricans maintain individual feeding territories. These territories are distributed contiguously, forming distinct colonies. Females visit male territories to spawn, and eggs are guarded by males until hatching. Male-male competition and female mate choice were studied in two colonies of different size compositions. Only larger individuals bred in both colonies. Some males in the large colony, that were larger than the breeding males in the small colony, did not succeed in reproducing probably because of severe attacks by the larger males while courting. However, females did not choose large size among breeding males. The most important male characteristic in female choice was the frequency of courtship displays in both colonies. Females in the large colony chose males mainly on the basis of the frequency of displays conducted in the females' territories, whereas females in the small colony chose males on the basis of the frequency of displays conducted in the males' territories. This difference may be a result of the difference in colony size. The distances between females' and males' territories were much greater in the large colony, and, because females cannot see courtship displays conducted in distant male territories, males in the large colony may have had to visit female territories frequently in order to conduct courtship near the females.     

Competence Pheromones in Bacteria

This article deals with general concepts concerning bacterial pheromones, including sex pheromones. The data on the structure and mode of action of competence pheromones in streptococci and bacilli are discussed, raising the question of the selective value of the process of genetic transformation for bacterial life activities and evolution.     

Sex Pheromones of C. elegans Males Prime the Female Reproductive System and Ameliorate the Effects of Heat Stress

Pheromones are secreted molecules that mediate animal communications. These olfactory signals can have substantial effects on physiology and likely play important roles in organismal survival in natural habitats. Here we show that a blend of two ascaroside pheromones produced by C. elegans males primes the female reproductive system in part by improving sperm guidance toward oocytes. Worms have different physiological responses to different ratios of the same two molecules, revealing an efficient mechanism for increasing coding potential of a limited repertoire of molecular signals. The endogenous function of the male sex pheromones has an important side benefit. It substantially ameliorates the detrimental effects of prolonged heat stress on hermaphrodite reproduction because it increases the effectiveness with which surviving gametes are used following stress. Hermaphroditic species are expected to lose female-specific traits in the course of evolution. Our results suggest that some of these traits could have serendipitous utility due to their ability to counter the effects of stress. We propose that this is a general mechanism by which some mating-related functions could be retained in hermaphroditic species, despite their expected decay.     

Natural Variation in the Strength and Direction of Male Mating Preferences for Female Pheromones in Drosophila melanogaster

Many animal species communicate using chemical signals. In Drosophila, cuticular hydrocarbons (CHCs) are involved in species and sexual identification, and have long been thought to act as stimulatory pheromones as well. However, a previous study reported that D. melanogaster males were more attracted to females that were lacking CHCs. This surprising result is consistent with several evolutionary hypotheses but is at odds with other work demonstrating that female CHCs are attractive to males. Here, we investigated natural variation in male preferences for female pheromones using transgenic flies that cannot produce CHCs. By perfuming females with CHCs and performing mate choice tests, we found that some male genotypes prefer females with pheromones, some have no apparent preference, and at least one male genotype prefers females without pheromones. This variation provides an excellent opportunity to further investigate the mechanistic causes and evolutionary implications of divergent pheromone preferences in D. melanogaster males.     

Estimating Survival for Elusive Juvenile Pond-Breeding Salamanders

Juvenile vital rates have important effects on population dynamics for many species, but this demographic is often difficult to locate and track. As such, we frequently lack reliable estimates of juvenile survival, which are necessary for accurately assessing population stability and potential management approaches to conserve biodiversity. We estimated survival rates for elusive juveniles of 3 species, the ringed salamander (Ambystoma annulatum), spotted salamander (A. maculatum), and small-mouthed salamander (A. texanum), using 2 approaches. First, we conducted an 11-month (2016–2017) mark-recapture study within semi-natural enclosures and used Bayesian Cormack-Jolly-Seber models to estimate survival and recapture probabilities. Second, we inferred the expected annual juvenile survival rate given published vital rates for pre-metamorphic and adult ambystomatids assuming stable population growth. For all 3 species, juvenile survival probabilities were constant across recapture occasions, whereas recapture probability estimates were time-dependent. Further, survival and recapture probabilities among study species were similar. Post-study sampling revealed that the initial study period median estimate of annual survival probability (0.39) underestimated the number of salamanders known alive at 11 months. We therefore appended approximately 1 year of opportunistic data, which produced a median annual survival probability of 0.50, encompassing salamanders that we knew to have been alive. Calculation from literature values suggested a mean annual terrestrial juvenile ambystomatid survival probability of 0.49. Similar results among our approaches indicated that juvenile survival estimates for the study species were robust and likely comparable to rates in nature. These estimates can now be confidently applied to research, monitoring, and management efforts for the study species and ecologically similar taxa. Our findings indicated that similarly robust vital rate estimates for subsets of ecologically and phylogenetically similar species can provide reasonable surrogate demographic information that can be used to reveal key factors influencing population viability for data-deficient species. © 2020 The Wildlife Society.     

Responses of Second-Instar Male Nymphs of Four Mealybug Species (Hemiptera: Pseudococcidae) to Conspecific and Heterospecific Female Sex Pheromones

The response of the late second-instar male nymphs of the mealybug species (Hemiptera, Pseudococcidae), Planococcus citri (Risso), Planococcus ficus (Signoret), Pseudococcus cryptus Hempel Nipaecoccus viridis (Newstead), to their conspecific and heterospecific female pheromone was studied. Males that exhibited the typical appearance of late-second-instar nymphs were tested. The male behavior was monitored soon after their exposure to the tested female sex pheromone in glass Petri dish arenas. Male nymph behavior toward the pheromone source was characterized based on their aggregation on the disks in the arena. Males of all four tested mealybug species were attracted to their conspecific female pheromone. By contrast, almost no interceptions of male nymphs with disks impregnated with a heterospecific female pheromone were observed. The mode of attraction of each of male nymphs of P. ficus, among most of the tested individuals (>80%), to the conspecific female sex pheromone, (S)-lavandulyl senecioate and or (S)-lavandulyl isovalerate, was the same as the mode of attraction latter on as adult. We suggest that by being attracted to the conspecific pheromone these males may direct themselves to a suitable pupation site near conspecific non-sibling mature females, thus preventing inbreeding. The repellency of heterospecific sex pheromone to males that are looking for a pupation site suggests that the latter try to avoid close contact with heterospecific females.     

Effects of Road Crossings on Stream and Streamside Salamanders

Abstract: Salamanders are important members of faunal communities in Appalachian Mountain (USA) streams, and the use of salamanders as bioindicators is increasing. Roads are a part of the modern landscape, but effects of roads on stream and streamside salamander diversity and abundance is not clear. We sampled streams in central West Virginia, USA, using quadrats placed along transects in the flowing channel, dry channel, and stream bank to assess salamander diversity, richness, and abundance during 2004. We used Akaike's Information Criterion for model selection at reach (i.e., above and below culverts) and stream scales. Salamander diversity and richness was affected by elevation, stream gradient, canopy cover, and the presence of roads. Overall, stream and riparian habitat quality was the most important factor affecting salamander richness. The presence of roads, stream gradient, and elevation received the most empirical support for predicting species' abundances. Roads benefited disturbance-tolerant species but negatively affected other species. Impacts of roads and culverts on habitat should be considered by federal and state transportation agencies and natural resources agencies during the planning process and addressed through mitigation efforts. Managers should install culverts that are as wide as the stream channel, at grade with the streambed, and dominated by rubble substrate to provide maximum benefit for salamanders. (JOURNAL OF WILDLIFE MANAGEMENT 72(3):760–771; 2008)