The origin of parental care in relation to male and female life history

The evolution of maternal, paternal, and bi‐parental care has been the focus of a great deal of research. Males and females vary in basic life‐history characteristics (e.g., stage‐specific mortality, maturation) in ways that are unrelated to parental investment. Surprisingly, few studies have examined the effect of this variation in male and female life history on the evolution of care. Here, we use a theoretical approach to determine the sex‐specific life‐history characteristics that give rise to the origin of paternal, maternal, or bi‐parental care from an ancestral state of no care. Females initially invest more into each egg than males. Despite this inherent difference between the sexes, paternal, maternal, and bi‐parental care are equally likely when males and females are otherwise similar. Thus, sex differences in initial zygotic investment do not explain the origin of one pattern of care over another. However, sex differences in adult mortality, egg maturation rate, and juvenile survival affect the pattern of care that will be most likely to evolve. Maternal care is more likely if female adult mortality is high, whereas paternal care is more likely if male adult mortality is high. These findings suggest that basic life‐history differences between the sexes can alone explain the origin of maternal, paternal, and bi‐parental care. As a result, the influence of life‐history characteristics should be considered as a baseline scenario in studies examining the origin of care.     

To hatch and hatch not: similar selective trade-offs but different responses to egg predators in two closely related, syntopic treefrogs

Risk-sensitive hatching is adaptive for species facing a trade-off between egg-stage and post-hatching risks, and environmental variation in one or both stages. Such plasticity has been found in amphibians, fishes, reptiles and spiders, with red-eyed treefrogs (Agalychnis callidryas) being the best-studied case. We assessed hatching plasticity and egg- and larval-stage risks in a closely related, syntopic species, the gliding leaf-frog (Agalychnis spurrelli). We found a lower hatching response to egg-eating snakes in A. spurrelli (9–28% of embryos escaped) than in A. callidryas (59–80% escaped). Levels of snake predation were similarly high for clutches of both species monitored at a pond in Costa Rica, and in fish predation experiments early-hatched A. spurrelli tadpoles were more vulnerable than later hatchlings, as has been shown for A. callidryas. A. spurrelli thus face a risk trade-off similar to A. callidryas, and likely would benefit from predator-induced hatching; their lower responsiveness to snakes appears nonadaptive. A. spurrelli embryos showed a stronger hatching response (57% hatched in 1 h) to submergence underwater than to snake attacks even though submergence is a less frequent risk. This suggests they have a greater capacity for early hatching than is expressed in the context of snake attacks, but have much lower sensitivity to snake cues than to flooding cues. Development in A. spurrelli is accelerated compared to syntopic A. callidryas, and spontaneous hatching is earlier and more synchronous. This is congruent with predictions based on selection by egg predators in the absence of a strong escape hatching response.     

Interactions between two closely related phytal harpacticoid copepods, asymmetric positive and negative effects

Competition for food is generally thought to exert a strong evolutionary pressure, driving trophic niche separation, either by specialization and/or by widening the choice of potential food resources. Harpacticoid copepods are common inhabitants of phytal assemblages, where several closely related species of the so-called phytal dwelling families often co-occur. However, direct competition among phytal harpacticoids has been thought to be unlikely, due to the abundant and continuously available food supplies. We conducted a series of field and laboratory studies to assess the role of competition in the abundance distribution of two closely related harpacticoid species, Mesochra rapiens and M. aestuari. We found that the abundance of both species co-varied on several seaweed species in the northern Baltic Sea, during a 3-month period. Stable isotope ratios in the green alga Cladophora glomerata field samples indicated different resource utilization of the two species, both in fresh and deteriorated C. glomerata, and in drifting algae. We tested in the laboratory if resource utilization was different between the species in sympatry and allopatry. We used enriched stable carbon isotope ratios (¹³C/¹²C) of the diatom Phaeodactylum tricornutum to trace the uptake in both species. Results from these experiments showed a much higher assimilation by M. aestuari in sympatry with M. rapiens, while the latter species showed a higher assimilation in allopatry. Our results show that while there is no apparent competition for resources between these two species in the field, there seems to be an asymmetric reaction when in sympatry and provided one single resource in the laboratory. We suggest that M. rapiens may facilitate assimilation by M. aestuari and discuss the mechanisms by which this may take place.     

Juvenile hormone titre and egg production in Tenebrio molitor infected by Hymenolepis diminuta: effect of male and/or female infection,male age and mating

Infection of Tenebrio molitor with Hymenolepis diminuta induces curtailment of female fertility. We examined ovulation and oviposition, and associated titres of juvenile hormone (JH), in relation to parasitism and mating. Oviposition was significantly increased in infected mated and virgin beetles by days 6 and 9 post-emergence. Ovulation was not changed by infection; by the end of the 18-day experiment, the total number of laid eggs was not significantly altered. On day 6, JH levels were significantly higher in virgin infected insects, compared to non-infected controls (236+/-37.7 and 107+/-9.62 pg/g wet weight). Oviposition increased after mating, but total eggs ovulated remained the same. JH levels were higher in mated females on days 12 and 18 post-emergence, for infected and control insects. Previous studies suggested that male reproductive potential might rise following infection, because uninfected females lay more eggs when mated to infected males. We tested whether this caused an increase in female JH. Males were mated on days 5 or 12, when significant changes in their reproductive physiology begin to be observed, and are maximal, respectively. However, male age was of greater significance in promoting JH levels in females (p=0.001), than infection status of either partner (p=0.33).     

Body temperature patterns in two syntopic elephant shrew species during winter

We measured body temperature (T_b) in free-ranging individuals of two species of elephant shrews, namely western rock elephant shrews (Elephantulus rupestris) and Cape rock elephant shrews (E. edwardii), during winter in a winter-rainfall region of western South Africa. These syntopic species have similar ecologies and morphologies and thus potential for large overlaps in diet and habitat use. Unexpectedly, they displayed different T_b patterns. Western rock elephant shrews were heterothermic, with all individuals decreasing T_b below 30 °C on at least 34% of nights. The level of heterothermy expressed was similar to other species traditionally defined as daily heterotherms and was inversely related to T_a, as is commonly seen in small heterothermic endotherms. In contrast, Cape rock elephant shrews rarely allowed their T_b to decrease below 30 °C. The level of heterothermy was similar to species traditionally defined as homeotherms and there was no relationship between the level of heterothermy expressed and T_a. In both species, the minimum daily T_b was recorded almost exclusively at night, often shortly before sunrise, although in some individuals minimum T_b occasionally occurred during the day. The interspecific variation in T_b patterns among Elephantulus species recorded to date reiterates the importance of ecological determinants of heterothermy that interact with factors such as body mass and phylogeny.     

A contribution to the functional morphology of the midgut gland in phalangiid harvestmen Gyas annulatus and Gyas titanus during their life cycle

The structure of the midgut gland and its changes in different seasons have been examined in the harvestmen Gyas annulatus and Gyas titanus (Arachnida: Opiliones: Phalangiidae). In both species, in the epithelium of the midgut gland two different types of cells are present: secretory and digestive ones. The secretory cells are characterized by plentiful rER and secretory granula. The digestive cells are characterized by an apical system of tubules. Both cells are connected by prominent specialized junctions. If a secretory cell is in contact with a digestive cell, rER cisterna are in close vicinity and parallel to these junctions. As found light- and electron microscopically and also histochemically, glycogen and lipids are stored in both cells. In both species, glycogen was seen to be used as energy compound during overwintering. At the end of their life, the digestive cells develop into excretory ones, containing metabolic wastes.     

Intra- and interspecific host discrimination in two closely related egg parasitoids

Intraspecific host discrimination is frequently found in solitary parasitoids, but interspecific host discrimination, where female parasitoids recognize hosts already parasitized by females of other species, is rare. This particular behaviour appears to be adaptive only under specific circumstances. In this paper, we quantified intraspecific host discrimination in Anaphes n. sp. (Hymenoptera: Mymaridae), an endoparasitoid of the eggs of Listronotus oregonensis (LeConte) (Coleoptera: Curculionidae) and interspecific host discrimination toward eggs parasitized by Anaphes sordidatus (Girault), a sympatric species competing for the same resource in similar habitats. To examine host discrimination, choice experiments were used where the females had to choose between different categories of eggs (unparasitized, parasitized by Anaphes n. sp. or A. sordidatus). Superparasitism and multiparasitism were avoided in experiments where the female had a choice between unparasitized hosts and hosts parasitized by the same female, by a conspecific or by a female A. sordidatus. When all hosts available were parasitized, conspecific superparasitism occurred more often than self-superparasitism or multiparasitism. These results indicated that females Anaphes n. sp. were capable of self-, conspecific and interspecific discrimination. Self-discrimination followed recognition of an external marking while interspecific discrimination occurred mostly after insertion of the ovipositor. Interspecific discrimination could result from the recent speciation of these species and could be associated with a genotypic discrimination. This behavior appears to be adaptive because of the competition for common hosts between the two parasitoid species.     

The potential for egg-guarding care in females of the damselfish,Dascyllus reticulatus,in the absence of uniparental male care

Male damselfish typically demonstrate uniparental egg-guarding care in nature. Potential plasticity in sexual behavior has recently been reported in various teleost fish. To examine behavioral plasticity in parental care, we conducted aquarium experiments to explore the potential for egg-guarding care in the female damselfish, Dascyllus reticulatus. After initial caretaking, males were removed from the mating nests, and cohabiting females frequently exhibited egg predation on the same day. However, we confirmed that females showed significantly decreased egg-predation frequencies on the following day and showed egg-caring behaviors. All experimental females guarded their eggs until they hatched. Females subsequently spawned eggs as females even after performing parental care behaviors, indicating no progression of sex change into males. Molecular analysis of select pituitary gland hormones indicated that egg-caring females and males showed high expression levels of prolactin, suggesting its involvement in the development of parental care behaviors. The cryptic possession of caretaking ability in females may be a tactical response to the need for temporary replacement of the care roles in cases where caretaking males are removed, for example, through predation, in damselfish species living in sexually cohabiting groups.     

华南地区3种具有不同入侵性的近缘植物对低温胁迫的敏感性

以五爪金龙(Ipomoea cairica (L.) Sweet,重度入侵植物)、裂叶牵牛(Ipomoea nil (L.) Choisy,轻度/非入侵植物)和三裂叶薯(Ipomoea triloba L.,非入侵植物)3种起源于热带美洲、且在华南地区具有不同入侵性的番薯属藤本植物作为研究对象,通过比较它们在低温胁迫下的生理响应探究3种植物对低温的敏感性与它们入侵性之间的关系。通过测定在不同温度(28、15、10 ℃)处理下植物的生物量、活性氧、渗透调节物质、根系活力、光合特征等生理指标发现,五爪金龙、裂叶牵牛及三裂叶薯均通过增加光合系统Ⅱ的热耗散、积累渗透调节物质以及增强根系活力来应对低温环境,但15 ℃的温度条件已经对3种植物形成较为强烈的胁迫作用,表现为H₂O₂和丙二醛的积累、光合系统Ⅱ受损、根部细胞死亡以及生物量、根长的极显著下降(P<0.01),证明3种植物对低温胁迫均具有较高的敏感性。综合比较3种植物各生理指标的响应幅度发现,它们对低温的耐受性表现为:五爪金龙 > 裂叶牵牛 > 三裂叶薯,这与它们在华南地区的入侵危害程度一致,暗示低温敏感性的差异可能是其入侵性差异的重要原因。结果表明,低温敏感性是影响外来植物入侵性和入侵区域的重要因素,五爪金龙较高的低温敏感性是限制其在华南以外地区形成入侵危害的重要原因。     

Female egg dumping and the effect of sex ratio on male egg carrying in a coreid bug

Phyllomorpha laciniata Vill (Heteroptera, Coreidae) is uniqueamong terrestrial insects in that females glue eggs on the backsof other conspecifics. Egg carrying byP. laciniatamales haspreviously been considered as paternal care. We explored femaleoviposition with respect to previous mating experience of femalesand tested whether sex ratio affects male egg-carrying. Thehypothesis that male egg-carrying is a form of paternal carepredicts that a male should always accept eggs after matingwith a female. However, if male egg-carrying is a form of postcopulatorymate guarding rather than paternal care, egg carrying shouldincrease in the presence of other males. When two couples wereplaced together, females laid eggs on the backs of all individualsenclosed, including the backs of other females. However, whena female was accompanied by 2 males, 22 out of 26 females ovipositedon their mating partner. Thus, sexual competition rather thanpaternity alone, affects a male's eagerness to carry eggs. However,even if males sometimes carry their own eggs, females lay eggson the backs of all conspecifics they can easily acquire. Thus,egg carrying in P. laciniata is partially voluntary and partiallythe result of female egg dumping     

The evolutionary puzzle of egg size,oxygenation and parental care in aquatic environments

Offspring fitness generally improves with increasing egg size. Yet, eggs of most aquatic organisms are small. A common but largely untested assumption is that larger embryos require more oxygen than they can acquire through diffusion via the egg surface, constraining egg size evolution. However, we found no detrimental effects of large egg size on embryo growth and survival under hypoxic conditions. We tested this in the broad-nosed pipefish, Syngnathus typhle, whose males provide extensive care (nourishment, osmoregulation and oxygenation) to their young in a brood pouch on their bodies. We took advantage of this species'' pronounced variation in egg size, correlating positively with female size, and tested the effect of hypoxia (40% dissolved oxygen) versus fully oxygenated (100%) water on embryo size and survival of large versus small eggs after 18 days of paternal brooding. Egg size did not affect embryo survival, regardless of O₂ treatment. While hypoxia affected embryo size negatively, both large and small eggs showed similar reductions in growth. Males in hypoxia ventilated more and males with large eggs swam more, but neither treatment affected their position in the water column. Overall, our results call into question the most common explanation for constrained egg size evolution in aquatic environments.     

The male and female perspective in the link between male infant care and mating behaviour in Barbary macaques

Infant care from adult males is unexpected in species with high paternity uncertainty. Still, males of several polygynandrous primates engage in frequent affiliative interactions with infants. Two non‐exclusive hypotheses link male infant care to male mating strategies. The paternal investment hypothesis views infant care as a male strategy to maximize the survival of sired offspring, while the mating effort hypothesis predicts that females reward males who cared for their infant by preferably mating with them. Both hypotheses predict a positive relationship between infant care and matings with a particular female. However, the paternal investment hypothesis predicts that increased matings come before infant care whereas the mating effort hypothesis predicts that infant care precedes an increase in matings. Both hypotheses are usually tested from the perspective of the proportion of matings and care that individual females engage in and receive, rather than from the perspective of the care and mating behaviour of individual males. We tested the relationships between care and mating from both female and male perspectives in Barbary macaques. Mating predicted subsequent care and care predicted subsequent mating when viewed from the male but not the female perspective. Males mainly cared for infants of their main mating partners, but infants were not mainly cared for by their likely father. Males mated more with the mothers of their favourite infants, but females did not mate more with the main caretakers of their infants. We suggest that females do not choose their mating partners based on previous infant care, increasing paternity confusion. Males might try to increase paternal investment by distributing the care according to their own instead of female mating history. Further, males pursue females for mating opportunities based on previous care.