Article

Paternity and the evolution of parental care

Authors:
To read the full-text of this research, you can request a copy directly from the authors.

Abstract

It is generally believed that level of paternity (the proportion of zygotes in a brood that were fertilized by the male providing parental care) has an important role in the evolution of parental care. We have used population genetics models to investigate this role. The models indicate that only in mating systems where a parental male “sacrifices” promiscous matings can paternity influence the evolution of male parental care. This is because level of paternity can reflect the number of opportunities for these promiscuous fertilizations. For example, high paternity can mean few opportunities and therefore a low cost for paternal care.Certain behaviors may preadapt a species for the evolution of male parental care because they decrease the costs of providing care. For example, in fish species where male care has evolved from spawning territories, the very establishment of territories may have precluded males from gaining promiscuous matings, thereby eliminating the promiscuity costs and facilitating the evolution of care. Without a promiscuity cost, level of paternity will not have influenced the evolution of male care in fishes.Because paternity has limited influence in the evolution of male care, differences in reliability of parentage between males and females are unlikely to explain the prevalence of female care. Our analysis suggests that paternity differences between species cannot serve as a general explanation for the observed patterns of parental care behavior.

No full-text available

Request Full-text Paper PDF

To read the full-text of this research,
you can request a copy directly from the authors.

... However, there are a relatively small number of species in which sires take care of their offspring together with females (Hartung and Dewsbury, 1979;Elwood, 1983;Dewsbury, 1985;Gromov, 2013a). This is seen as an evolutionary tradeoff, which, on the one hand, improves the survival conditions of the young, and, on the other hand, limits the freedom of male mating and adversely affects their competitiveness (Trivers, 1972;Werren et al., 1980;Dewsbury, 1985). Care for offspring is demonstrated primarily by males of those species that are characterized by a family-group lifestyle (Elwood, 1975;Hartung and Dewsbury, 1979;McGuire and Novak, 1984;Gromov, 2013a). ...
... The generally accepted concept of the evolution of paternal behavior prescribes the existence of an obligatory connection between the demonstration of parental care in males and the increase in their individual fitness (Kleiman, 1977;Werren et al., 1980;Wittenberger and Tilson, 1980). The idea of individual fitness is the key in socioecological concepts (Crook et al., 1976, Lacey andSherman, 2007), which explain the evolution of sociality (group way of life) in mammals, including rodents. ...
Article
Pup survival and growth rates were monitored to determine whether paternal care potentially contributes to reproductive success of male common voles Microtus arvalis Pallas, 1779. The subjects were 258 pups from 47 litters obtained from adult voles trapped in the wild. Reproducing pairs were formed and divided onto two groups in accordance with the age and parental experience of the reproducing individuals: group I included pairs with subadult males and nulliparous females; group II included pairs with adult males and multiparous females. Each of the groups was divided into two subgroups: in subgroups Iа (n = 13) and IIа (n= 11), litters were raised by females without sires; in subgroups Ib (n = 12) and IIb (n = 11), litters were raised by both parents. The litter size was recorded at parturition and again when the young were 12, 21, and 30 days old. Pups in each litter were weighed at parturition and also again when the young were 12, 21, and 30 days old. This study has shown that the presence of the sire may negatively affect both survival and growth rates of the pups, but this effect is dependent on the age of the reproducing individuals.
... Существует мнение, что формированию прочных парных связей и, как следствие, проявлению отцовской заботы о потомстве способствует низкая плотность населения, при которой энергетические затраты на поиск рецептивных самок существенно возрастают (Maynard-Smith, 1977;Carter, Getz, 1993;Getz, Carter, 1996). Многие исследователи рассматривают отцовское поведение у грызунов как эволюционный компромисс (trade-off), обеспечивающий улучшение условий выживания относительно немногочисленного потомства за счет ограничения возможности свободного спаривания самцов с наибольшим числом самок и, соответственно, более широкого распространения своих генов в популяции (Trivers, 1972;Werren et al., 1980;Dewsbury, 1985). ...
... Иными словами, охраняя территорию, самцы проявляли косвенную заботу о потомстве, которое лишь отчасти являлось их собственным (McLean, 1983). Этот факт согласуется с предположениями некоторых ученых-теоретиков, пола-гающих, что отцовская забота и промискуитет не относятся к взаимоисключающим явлениям (Werren et al., 1980). Об этом свидетельствуют также и прямые наблюдениями за видами, ведущими семейно-групповой образ жизни, с которым беспорядочные спаривания, казалось бы, несовместимы (Громов, 2008). ...
Book
В монографии обобщены результаты многолетних полевых и лабораторных исследований автора, проведенных в 1976-2012 гг. и дополненных литературными данными о поведении более 40 видов грызунов. Рассмотрены физиологические и этологические механизмы формирования и регуляции родительского поведения, а также его связь с пространственно-этологической структурой популяций грызунов и формированием феномена социальности в эволюционном аспекте
... An alternative male reproductive strategy, that is, a display of caring for offspring, is indicated in a small number of species (Kleiman, 1977). Paternal behavior in this case is considered an evolutionary compromise (trade-off), improving the conditions of offspring survival but limiting the freedom of mating and the competitive ability of males (Trivers, 1972;Werren et al., 1980). Parental behavior is mostly expressed in species with strong pair bonds and a family-group way of life (Hartung and Dewsbury, 1979;McGuire and Novak, 1984;Gromov, 2013а). ...
... The advantages related with offspring rearing in these species are reckoned to prevail over the possible losses of males related with the limitation of mating freedom (Gubernick et al., 1993). This assumption is laid into the base of the male-care hypothesis, explaining the evolution of the parental behavior of males (Kleiman, 1977;Werren et al., 1980;Wittenberger and Tilson, 1980), which expressed that male participation in rearing pups should ensure better offspring survival and thereby improve reproductive success. This hypothesis was subject to repeated verification, particularly to ascertain whether the presence of sire can contribute to the survival and growth rates of pups (Dudley, 1974;Elwood and Broom, 1978;Gerling and Yahr, 1979;Gubernick et al., 1993;Wynne-Edwards and Lisk, 1989). ...
Article
Two groups of litters obtained from individuals captured in natural environments have been monitored under laboratory conditions. The females of one group reared the offspring in the absence of sires; the litters of the other group are raised by both parents. It is ascertained that the presence of sires, especially those who are characterized by higher levels of aggression, has a negative effect on the survival and growth rates of the pups. Therefore, the reproductive success of common vole males can decrease if they enter the family-group composition and depends on the specificity of their behavior, first and foremost, an inclination toward exhibiting aggression.
... However, the low encounter rates and long inter-mating intervals of males in the field suggest that it is not very costly for males to remain with one female for a long period of time. Males should increase their paternity from a given mating only if in doing so they increase their net reproductive success (Parker 1970a, b; Werren et al. 1980). As Werren et al. (1980) pointed out, increasing investment in one female or her offspring does not necessarily cost males opportunities to mate with additional females. ...
... Males should increase their paternity from a given mating only if in doing so they increase their net reproductive success (Parker 1970a, b; Werren et al. 1980). As Werren et al. (1980) pointed out, increasing investment in one female or her offspring does not necessarily cost males opportunities to mate with additional females. Because of low densities and low encounter rates, males of L.c. clivicollis may not suffer a loss of opportunities to inseminate additional females by engaging in prolonged mating. ...
Article
Full-text available
Males and females of the milkweed leaf beetle mate for up to 2.5 d in the field; copulation is interspersed with mounted courtship behavior and periods of passive riding on the female's back. In 1985, females mated with up to 10 different males in Bridgeport, New York, and matings averaged 0.75-I-0.04 (SE) d in duration. Sperm utilization patterns are complex in the milkweed leaf beetle. In the laboratory, mating duration, mating order and the duration of the gap between matings affected paternity. The second male's sperm predominated when a female's two consecutive matings were of equal duration (but were long: 45 h each) or when they were intermediate in duration (15 h each) with a 5 day gap between the two matings. The mechanism of sperm predominance appears to be one of time-dependent sperm removal and replacement. Neither relative nor absolute size of males affected the extent to which they superceded the sperm of rivals. Electrophoretic analysis of paternity at 6 polymorphic loci showed that offspring of at least eight out of 11 mating pairs pairs collected in the field at the end of the 1984 breeding season were sired by more than one male. At least seven of the 11 families produced some offspring that were sired by the female's most recent mate in the field and in no case was the collected male completely excluded from paternity. In a greenhouse plot, males climbed off females' backs a mean of 1.1 _+ 0.3 h before oviposition. Females did not oviposit with males on their backs and did not oviposit sooner when males were removed than when males were allowed to remain mounted, indicating that males don't keep females from ovipositing until they can achieve sperm predominance. * Present address." Hastings Natural History Reservation, University of California, Star Route Box 80, Carmel Valley, CA 93924, USA The close concordance between male disembarkment and oviposition suggests that males remain with females beyond the time when they have superceded the sperm of their prior mates, possibly to guard them from rival males. Although the prolonged mating association of the milkweed leaf beetle cannot be separated into discrete copulatory and guarding phases, the results of this study suggest that prolonged mating enables males both to remove and replace the sperm of the females' prior mates with their own sperm and to prevent females from remating prior to oviposition.
... Smith (1978) Yes Paternity does not matter Grafen (1980) Yes Paternity does not matter Werren et al. (1980) Not necessary Paternity matters if paternal care trades off with EP fitness Winkler (1987) Yes Paternity always matters Whittingham et al. (1992) Yes Paternity always matters Xia (1992) Yes Paternity always matters Yamamura and Tsuji (1993) Yes Paternity always matters Westneat and Sherman (1993) Yes Paternity matters sometimes Houston (1995) Yes Paternity always matters Queller (1997) Yes Paternity always matters Kokko (1999) Yes Paternity always matters Houston and McNamara (2002) Yes Paternity always matters Fromhage et al. (2007) Yes Paternity always matters Kokko and Jennions (2008) Yes Paternity always matters Fromhage and Jennions (2016) Yes Paternity always matters Requena and Alonzo (2017) Yes Paternity always matters opportunities to increase his paternity by preventing his mate from mating with competitors. Hence, we find that mechanisms such as paternity guarding can lead to greater selection for male care even under conditions of partial paternity and complete maternity. ...
Article
The theoretical literature predicts that parentage differences between the sexes, due to females mating with multiple males, select males to provide less parental care and females to care more for the offspring. We formulate simple evolutionary games to question the generality of this prediction. We find that the relationship between paternal care and fitness gained from extra-pair matings is important. A trade-off between these two quantities is required for partial paternity and complete maternity to bias the ESS towards more female care. We argue that this trade-off has been implicitly or explicitly assumed in most previous theories. However, if there is no trade-off between paternal care and extra-pair matings, parentage differences do not influence the ESS sex roles. Moreover, it is also possible for these two quantities to have a positive relationship, in which case we predict selection for male care is possible. We support these predictions using agent based simulations. We also consider the possibility that caring males have greater opportunities to guard their paternity, and find that this mechanism can also select for male biased care. Hence we derive the conditions under which male care may be selected despite partial paternity and complete maternity. This article is protected by copyright. All rights reserved
... Maynard Smith (1977) and Werren et al (1980) claim that paternity certainty is only an issue in the evolution of paternal care if care giving is costly to the male in terms of missed mating opportunities. For example, in fish, males who establish spawning territories actually have very few opportunities for promiscuous mating and hence should provide care regardless of paternity (Werren et a i 1980). ...
Thesis
Humans are social animals. Human societies emerge from vast networks of cooperative interactions between many different individuals. In this respect, humans are similar to most other primates. However, human societies are unusual among primates in the number of different types of cooperative relationships that are involved. In humans, males and females form strong pair bonds within large multimale, multi-female societies in which many other cooperative relationships are also important. How and when did human social systems arise? Do males and females use different types of cooperative strategies? Under what conditions does paternal care evolve? Do males and females have different constraints, and how do these affect the types of social strategies they employ? How do factors such as environment quality and seasonality modify these strategies? This thesis seeks answers to these questions using computer simulations based on the iterated Prisoner's Dilemma. The hypotheses generated by these models are tested using data from living primates. They are then used to investigate the kinds of societies that our hominid ancestors may have lived in. The theoretical and empirical evidence presented in this thesis suggests that sex differences in the energetic cost of reproduction determine the cooperative strategies, and ultimately the types of social groups, that evolve. It is proposed that during hominid evolution female energetic costs increased greatly, in comparison to male energetic costs, due to changes in body size dimorphism, diet and brain size. A two-stage model of hominid social structure is developed. The first stage, at the transition from the australopithecines to Homo erectus, would have involved an increase in female cooperation, especially food sharing. The second stage, occurring between 500,000 and 100,000 years ago, would have involved male care giving, the formation of pair bonds and the sexual division of labour within the context of a wider cooperative network.
... The prevailing paradigm assumes that a male's fitness can increase through providing care if his offspring survive and reproduce, and certainty of paternity is presumably a contributor to the evolution of paternal care. However, certainty of paternity is shown to be not required for paternal care in rodents (Hartung & Dewsbury, 1979;Werren et al., 1980). Therefore, male fitness benefits do not adequately explain the evolution of paternal care among rodents. ...
Article
Full-text available
The evolution of paternal care in rodents has intrigued biologists for over decades. In this paper, both ultimate (adaptive significance, evolution) and proximate (ontogeny, mechanisms) questions related to the emergence and maintenance of male paternal care are reviewed. Paternal care is thought to be a consequence of social monogamy, but no definitive hypothesis adequately explains the evolution of paternal behavior in rodents. The onset, activation and maintenance of paternal care are shown to be governed by complex interactions in neuroendocrine systems that change during ontogeny. Depending on the species, different components of male experience as well as different exogenous cues are likely to be involved in the organization and activation of paternal behavior. Several hormones, including steroids (testosterone, estradiol, progesterone) and neuropeptides (prolactin, vasopressin, oxytocin), are involved in the onset, the maintenance, or both the onset and the maintenance of parental behavior, including direct paternal care. The effect of testosterone was found to be not universal and, moreover, species-specific. As for estrogens and neuropeptides, further investigations are needed for a better understanding of the role of these hormones in activation and maintenance of rodent paternal behavior. Current research shows that male parental care in rodents is, to a great extent, an epigenetic phenomenon, and future studies will focus on the epigenetic modifications that can affect the paternal behavior in rodents.
... Uma vez que uma maior quantidade (e maiores distâncias) nos deslocamentos foram observadas no inverno, é provável que os mesmos sejam realizados tanto para busca de recursos alimentares quanto para busca de parceiros. Na região Neotropical, o período mais quente e chuvoso é associado à maior disponibilidade de recursos (Lowe-McConnel, 1999 (Werren et al., 1980;Gross & Sargent, 1985), porém os resultados deste trabalho também não fornecem evidências para tal conclusão. ...
Thesis
Full-text available
Coastal streams from southeastern Brazil, due to the presence of the Serra do Mar ridge, are subject to violent short-term floods that can occur when heavy rains reach the headwaters (i.e. flash floods). These phenomena, which are more common during the rainy season, can also occur at any time during orographic rainfall. They are unpredictable and can also make streams unstable. This study aimed to verify the influence of this instability on populational characteristics, reproductive cicles, life history and possible movement patterns of the catfish Rhamdioglanis transfasciatus, an endemic species from the Atlantic Rainforest. Fish sampling were carried out in the Guapiaçu river basin (RJ), using electric fishing. For the reproductive study, individuals were captured, anesthetized, euthanized, measured, weighed and dissected for analysis and weighing of gonads, stomach and liver. Individuals used in the species movement patterns study were captured, anesthetized, measured, marked subcutaneously with visible implants (AlphaTag VI®) and returned to the midpoint of the stream stretch from which they were originaly captured. R. transfasciatus presented a long reproductive period, starting at the beginning of winter and remaining until spring. The triggers for the onset of reproductive activity in females and males are linked respectively to factors of greater temporal and spatial scale (decrease of rainfall and decrease of water temperature) and with more local factors (decrease in rainfall accumulated in recent days and decrease in stream flow). The females' fecundity was relatively low, but increased as there was biomass gain. We found evidence for sexual dimorphism in body size, caudal fin size and body height. The sexes have different growth patterns and size at first maturity, which are probably associated with divergent energetic demands. During the cold and dry period (winter), when the risk of flash floods is lower and the species is also reproductively active, individuals of R. transfasciatus showed extended home ranges. The proportions and orientation of movements are probably linked to the ease of movements in favor of flow. There was a decrease in the proportion of growth of R. transfasciatus that displaced longer distances, which suggests higher energy expenditure. The low individual reproductive investment associated with a long reproductive period suggests non synchronous reproductive activity among individuals, which can be interpreted as being adaptive to an unstable environment. The role of displacements in the reproductive period is still unknown, but it can be assumed that it is a mechanism of reduction of intraspecific competition in the use of space and the obtaining of food. The energetic relationships linked to the reproduction and growth of R. transfasciatus and other stream fish species, subjected to environmental instability, still need further investigation.
... Dolayısıyla eğitim konusunda toplumsal açıdan bir bilinçlenme sağlandığı söylenebilmektedir.Babalık Rolü ve Çocuk BakımıÇocuk bakımında annelik rolü kadar babalık rolü de önemlidir. Bu bağlamda erkeklerin sahip oldukları önemli rollerden biri de babalık rolüdür (Werrren, Gross veShine, 1980). Kadın katılımcılar tarafından babalık tanımlamaları farklılaşmaktadır. ...
Article
Full-text available
Ataerkillik ve geleneksel toplum yapısı erken yaşta evlilikleri meşru hale getiren bir zihniyetin ürünüdür. Bu zihniyetin bir ürünü olan erken evliliklerle birlikte kadın-erkek arasında eşit olmayan ilişkiler ortaya çıkmakta ve kadına yönelik bakış açısından kaynaklı bu eşitsizlik daha da artmaktadır. Özellikle erken evliliklere erkek çocuklarına nazaran daha çok kız çocuklarının maruz kalması bu durumun en önemli göstergelerinden biridir. Araştırma, erken evlilik yapan kadınların toplumsal cinsiyet algısını ele almaktadır. Araştırmanın amacı, 18 yaşından küçük evlilik gerçekleştiren kadınların ifadelerinden hareketle erken evliliğe ilişkin toplumsal cinsiyet rollerinin nasıl kurgulandığını, evliliklerinde yaşadıkları zorlukları, evlilik beklentilerini, rol paylaşımı sırasında yaşanan toplumsal cinsiyet eşitsizliğinin neler olduğunu diğer bir deyişle toplumsal cinsiyet algısını ortaya koymaktır. Buradan hareketle araştırma, Ağrı ilinde 15 kadın katılımcı ile yarı-yapılandırılmış bir görüşme formu kullanılarak ve derinlemesine görüşmeler yapılarak gerçekleştirilmiştir. Bu çerçevede araştırma erken evlilik yapan kadınlara uygulanarak onların toplumsal cinsiyet rollerini ele alması bakımından önem taşımaktadır. Araştırma sonucunda erken evliliklerle birlikte kadına yönelik toplumsal cinsiyet eşitsizliği daha da artmakta ve cinsiyet rolleri bakımından ikincil konuma düşmektedir.
... The result of this comparative study does not invalidate the present function ofcourtship feeding as paternal investment in bushcrickets. Males with large investments should have a high confidence of paternity (Werren et al. 1980;Gwynne 1984b), which assumes that the donating males will father the benefitted offspring (Gwynne I988b). A paternal investment function may have arisen secondarily, once paternity assurance is high. ...
Article
During courtship and copulation, males of many insect species provide the female with a nuptial gift of a prey item or synthesized material. These gifts may be explained as a form of paternal investment by increasing female reproductive output, or in terms of mating effort by increasing male fertilization success. These explanations, while not mutually exclusive, are controversial. While experimental studies examine the maintenance of nuptial gifts in single species, comparative studies are required to indicate more general evolutionary trends. Male bushcrickets provide females with a nuptial gift, a spermatophylax, which is transferred to females at mating along with the sperm-containing ampulla. Analysis of comparative data of 28 species of bushcrickets (Orthoptera: Tettigoniidae), reveals that male spermatophore size (spermatophylax and ampulla weight) is positively correlated with female refractory period, which, in turn, correlates with male fertilization success. Moreover, gift size (the spermatophylax) covaries with ejaculate size (the ampulla), which is consistent with the hypothesis that it serves as a sperm protection device. In contrast, there is no significant correlation between any measure of female fecundity and male spermatophylax size. This indicates that the variation in spermatophore size among bushcrickets is better explained by a mating-effort function than a paternal investment function.
... Defence of a breeding territory will maximise the chance that the brood being guarded is genetically related to the care giving parent. Werren et al. (1980) concluded that the level of paternity can only influence the evolution of male parental care where the male reduces his genetic costs by sacrificing the opportunity for promiscuous matings. Baylis (1978Baylis ( , 1981 argues that it is the relative rate rather than the energetic cost of production of gametes that determines the sex of the care-giving parent. ...
Thesis
Full-text available
The Gadopsidae is an endemic mono-generic freshwater fish family of south-eastern Australia. The family was thought to be mono-specific, containing the River Blackfish Gadopsis marmoratus, until 1984 when a second species, the Two-spined Blackfish G. bispinosus was described (Sanger 1984). Due its relatively recent description, little information was available on the ecology or conservation status of G. bispinosus in NSW and the ACT. The distribution and abundance of G. bispinosus in southern NSW and the ACT was investigated with the species recorded at 16 of 119 sites surveyed. At most sites where they were recorded, G. bispinosus was abundant, and was invariably found in association with one of the trout species. G. bispinosus was found to be restricted to cool, clear upland streams with rocky, cobble bottoms and relatively intact forest vegetation. It was hypothesised that the presence of the species in the upper Murrumbidgee drainage was due to stream capture, in which the headwater streams from the Murray drainage have been captured by the Murrumbidgee drainage. Interpretation of historical reports of blackfish distribution in NSW indicated that G. bispinosus had suffered declines in some rivers with the species now apparently absent from the Yarrangobilly River and the Murrumbidgee, Naas, and Paddys rivers in the ACT. The decline is thought to have been due to habitat degradation, particularly sediment addition, which has reduced the cover available by filling the interstitial spaces in the cobble substrate favoured by G. bispinosus. The invariable association of G. bispinosus with introduced trout species and habitats with abundant cover, suggests that trout may have played some part in the current distribution of G bispinosus by excluding them from sub-optimal habitats. The movements of G. bispinosus was found to be very restricted with a home range of approximately 15 metres estimated for adult fish. Recapture rates were high, particularly in adult fish, in comparison with other studies of freshwater fish, indicating that adult G. bispinosus are particularly sedentary. The home ranges of G. bispinosus were found to be stable from year to year with fish able to maintain their position in the stream over the high flow periods of winter and spring. On the basis of aquarium observations of pugnacious and aggressive behaviour between adult G. bispinosus, it was considered that these home ranges may be considered territories under the definition of (Gerking 1953) who defined a territory as "any defended area". The diet of G. bispinosus was investigated with distinct seasonal and ontogenetic differences apparent in the diet. Juvenile fish consumed predominantly smaller items such as early instar ephemeropterans and chironomid larvae with some trichopterans present in the diet. The proportion of trichopterans in the diet increased with increasing fish size, with the importance of ephemeropterans and dipterans inversely related to fish size. terrestrial items were not present at all in the stomachs of juvenile fish, were of minor importance to immature fish, but were a major dietary item of adults. Terrestrial items were most abundant in the diet of both immature and adult fish in summer and autumn. The diet of Rainbow Trout 0. mykiss was also examined, with significant dietary overlap apparent between 0. mykiss and G. bispinosus. As with G. bispinosus, the diet of 0. mykiss was dominated by ephemeropterans, trichopterans and terrestrial items. Dietary overlap was greatest between similar size classes of both species, with some seasonal pattern evident. Significant overlap occurred between the non-adult G. bispinosus and juvenile 0. mykiss all seasons except autumn. Overlap was greatest between mature blackfish and non-juvenile trout with significant dietary overlap recorded in all seasons. It was considered that the consistent significant overlap values indicate the dietary competition is likely between these two fish species. The reproductive ecology of G. bispinosus was found to be similar to that recorded for G. marmoratus by Jackson (1978a). Both blackfish species deposit large, yolky, adhesive, demersal eggs in late spring/early summer when water temperatures exceed 16-17 °C. Fecundity in G. bispinosus is low with less than 300 eggs carried by most females. The natural spawning site was not located but is thought to be in the interstitial spaces between cobble and boulders on the river bed. Artificial P.V.C. spawning tubes proved successful with a total of 15 egg masses deposited in them over the course of the study. The numbers of eggs in each egg mass were within the fecundity estimates of individual fish, and all eggs within a mass were at the same stage of development , indicating that a single fish is probably responsible for each egg mass. A large adult male was present with each egg mass. Eggs hatched after approximately 15-17 days, with the embryo emerging from the chorion but the yolk sac remaining inside, effectively tethering the young to the spawning substrate. Parental care by the male continued for approximately 3-4 weeks after hatching by which time the yolk sac was almost fully utilised and young blackfish could swim well.
... Indeed, peripheral blockade of kappa-opioid receptors decreased selective aggression in male prairie voles identifying the importance of this system in aversive social motivation. Consistent with previous studies, males were more aggressive than females and higher levels of aggression in males may be associated with uncertain paternity (Werren et al., 1980). Indeed, if males do not engage in mate guarding, some paired female prairie voles will mate with novel males . ...
Article
The socially monogamous prairie vole is an excellent animal model for studies of the neurobiology of selective social attachment. Here, we have demonstrated that monogamous bonds are formed and maintained by a balance between mu- and kappa-opioid receptor activation within motivational circuitry. We first show that the formation of a pair bond requires the activation of mu-opioid receptors that mediate both motivational and positive hedonic aspects of reward processing. We hypothesize that these two populations of mu-opioid receptors work in parallel to motivate social interactions with a potential mating partner and subsequently encode these interactions as rewarding through the induction of a positive hedonic state. In contrast to pair bond formation, which is associated with affiliative social encounters generally categorized as positive, the maintenance of a pair bond is associated with aversive social encounters, such as the aggressive rejection of novel conspecifics, that acts to both guard the mate and prevent the formation of a subsequent alternate bond. In the present body of work, we demonstrate that selective aggression, an established behavioral indicator of pair bond maintenance, is mediated by interactions between D1-like dopamine receptors and kappa-opioid receptors within the nucleus accumbens shell and that direct activation of kappa-opioid receptors within this region encodes social aversion. These data suggest that the encoding of social stimuli besides the mating partner as aversive and subsequently aggressively rejecting this stimulus is important for the maintenance of the original pair bond. Additionally, we show that the establishment of a pair bond is associated with a dramatic reorganization of the dopamine and kappa-opioid receptor systems, which likely acts to mediate the transition from generally affiliative to selectively aggressive. Together, these data suggest that interactions between valence coding opioid receptors and motivational circuitry are critical for guiding the direction of socially motivated behaviors, such as the motivation to form and maintain bonds. Importantly, an increased understanding of neural mechanisms that mediate social attachment behavior of prairie voles may provide insight into neural mechanisms that regulate attachment behavior in our own species.
... The second argument was criticized extensively by Maynard Smith (1978) and Werren et al. (1980), who argued that, assuming paternity remained constant across broods, uncertainty of paternity should not influence the evolution of male care. This argument was in turn criticized as incomplete (e.g., Westneat and Sherman, 1993). ...
Article
Tropical frogs exhibit an extraordinary diversity of reproductive strategies, including complex patterns of parental care and mating behavior. The neotropical poison frogs (families Dendrobatidae and Aromobatidae) have parental care in almost all species, and many different types of parental care have evolved, including male, female, and biparental care. These different patterns of care are associated with various aspects of ecology and life history, and with diverse mating strategies and systems. We discuss the ecology and evolution of parental care in the neotropical poison frogs in an evolutionary and ecological context, and discuss how sexual selection affects and is affected by parental care in this clade. We also review theoretical and empirical research on the evolution of parental care, sexual selection, and mating systems. We describe the patterns of parental care and sexual selection in poison frogs in the context of previous theoretical and empirical studies. Finally, we suggest potentially productive directions for future research.
... An inverse relationship between paternity and male parental care has been suggested by some theoretical models (Houston and Davies, 1985;Houston, 1995;Kokko, 1999;Ridley, 1978;Trivers, 1972;Westneat and Sherman, 1993;Whittingham et al., 1992;Winkler, 1987;Xia, 1992), but other models with different assumptions have not predicted such a cost of lost male parental care for females when sperm competition is prevalent (Grafen, 1980;Houston, 1995;Houston and Davies, 1985;Maynard Smith, 1977;Werren et al., 1980;Whittingham et al., 1992;Wittenberger, 1979). If certainty of paternity is the same in all breeding attempts of a male, then paternity should have no effect on optimal paternal behavior (Grafen, 1980;Maynard Smith, 1978). ...
Article
Full-text available
Paternity has been hypothesized to be related to the evolution of paternal care because (1) there should be selection for males not to invest in broods with an uncertain parentage, or (2) male extrapair activity is traded against paternal care. We used interspecific comparisons to discriminate between these alternatives. Male participation in three kinds of parental care (nest building, incubation, provisioning of offspring) increased with high paternity in their own nests. Male parental activities at some stages of the breeding cycle were significantly correlated. A multivariate analysis taking this intercorrelation between different components of care and potentially confounding variables such as precociality, polyandry, and sexual dichromatism into account revealed that paternity was significantly positively related to offspring provisioning, while male participation in the other components of parental care did not explain a significant amount of interspecific variation in paternity. Analyses of evolutionary transitions between different dichotomized states of paternity and paternal care provided no clear conclusions concerning evolutionary scenarios. However, theoretical arguments and the results of the contrast analyses suggest that male provisioning of offspring evolved in response to paternity.
... To understand the evolution of parental care, it is important to understand how reproductive success varies as a function of parental care. Parental care effort varies as individuals trade off between investment in the present offspring and future reproductive potential, with confidence in paternity (Werren et al. 1980, Winkler 1987, Whittingham et al. 1992, Xia 1992, Westneat and Sherman 1993, Houston 1995, Mauck and Grubb 1995, and with different mating systems (Maynard Smith 1977). ...
Article
Full-text available
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.
... The evolution of parental care in poison frogs has been established through decades of field observations and analysis of genetic markers [26,29,52]. Male uniparental care is the ancestral state in poison frogs (Figure 1d), and presumably evolved as a method of ensuring paternity [30,53]. The associated cost in time and resources by males caring for offspring likely also favored clutch size reduction and an increase in egg size [54]. ...
... Furthermore, in certain circumstances, females could benefit by cuckolding males into helping rear her infants and, to induce paternal responses, females might employ the normally mutually beneficial mechanisms that initiate a paternal state. This would be to the male's disadvantage if, by giving time to rear non-kin, he were to lose mating opportunities with other females (Werren et al., 1980). ...
Article
Parents may recognise and direct parental care towards their own offspring using one or more of several mechanisms. This review, however, focuses on situations where infants are encountered for the first time, when familarity cannot account for the protection of kin. Possible mechanisms by which kin may be protected are use of a genetic marker, association, location, and state-mediated recognition. There are some comments on female strategies to cuckold males, ie maternal kin protection and paternal non-kin protection. -P.J.Jarvis
... Paternal California mice appear to invest extensively in the care of their young. The evolution of paternal investment is often associated with monogamy (Kleiman, 1977;Kleiman & Malcolm, 1981;Wittenberger & Tilson, 1980), production of altricial offspring (Zeveloff & Boyce, 1980), low reproductive potential and occupying a stable habitat (Eisenberg, 1966;Kleiman, 1977), certainty of paternity (Alexander, 1974;Dawkins & Carlisle, 1976;Maynard Smith, 1978;Trivers, 1972; but see Werren, Gross, & Shine, 1980), and monopolization of resources and mates (Barash, 1975(Barash, , 1976Emlen & Oring, 1977;Orians, 1969). Our observations of extensive paternal behavior in Peromyscus californicus are consistent with the limited information available on this species in its natural habitat. ...
Article
Full-text available
Most knowledge of parent-offspring relations in mammals is derived from studies of mother-infant interactions. Male parental care has been less well studied. We explored maternal and paternal behavior of the California mouse, Peromyscus californicus. Six pairs of parents and their young were videotaped continuously for 12 hours/day, on alternate days from Days 1 to 31 postpartum. Males exhibit all parental activities and to the same extent as displayed by mothers, except lactation. Male parental behavior begins on the day of birth. Mothers and fathers spend substantial and equivalent amounts of time in the nest and in physical contact with pups throughout lactation. Males devote more time than females to licking pups, although females engage in more pup anogenital licking. Mothers nurse for at least 4 weeks, and fathers and mothers both build nests and carry young. The biparental care system of Peromyscus californicus affords an opportunity to develop a broader, more complete view of parent-offspring relations.
... F undamental to theoretical treatments of parental care evolution-and to many sexual selection and mating systems models-is the assumption that parental care is costly, specifically, that a trade-off exists between providing care and engaging in alternative current or future reproductive activities. The ways in which parental care is envisioned as costly vary, but in some models, providing care and seeking additional mates are viewed as mutually exclusive, such that the main cost of care is the number of extra offspring that could be produced during the parental care period if care was not pro vided (Maynard Smith 1977 ;Werren et al. 1980). The reproductive cycle is envisioned as divisible between "time-in," when the individual is available for mating, and "time-out," when it is engaged in parenting, replenishing gametes, etc., and is not available for mating (Clutton-Brock and Parker 1992;Kvarnemo and Ahnesjo 1996;Parker and Simmons 1996;Reynolds 1996;Kokko and Jennions 2008). ...
Article
Full-text available
Among avian species with biparental care, male alternative reproductive opportunities can occur in the form of either extrapair fertilizations or additional social mates. We manipulated testosterone (T) levels in male house sparrows to assess whether participation in parental care detracts from male success in securing alternative matings; we also compared the annual reproductive success of males that engaged in normal levels of care with the success of males that displayed increased mating effort at the expense of parental care. Our results showed that the incidence of polygyny among high-T/low-parenting males was elevated relative to control males, but that success in obtaining extrapair mates was independent of hormonal treatment. Thus, male parental care seems to carry an opportunity cost in terms of reduced acquisition of additional social mates. Despite this cost, between-treatment comparisons of the estimated number of fledglings sired annually suggest that, in this species, the value of male contributions to care is sufficiently high to favor the reductions in T that facilitate normal male parenting.
... Blumer (1982) considered that 89 families of fishes perform some kind of parental care, and in the majority of DOI: 10.1590 these species this role of a caregiver is played by the male. Trivers (1972) suggested that paternal investment in parental care is directly related to the certainty of paternity, that is, the more the male is in doubt of its paternity, the less will be its willingness to invest in the parental care, although this concept is not universally accepted (Werren et al., 1980;Bouwman et al., 2005). A number of studies have shown this phenomenon in birds (e.g., Moller & Cuervo, 2000;Peterson et al., 2001), fishes (e.g., Neff, 2003;Neff & Gross, 2001), and invertebrates (Hunt & Simmons, 2002). ...
Article
Full-text available
Monogamy is rare in fishes and is usually associated with elaborate parental care. When parental care is present in fishes, it is usually the male that is responsible, and it is believed that there is a relationship between the high energetic investment and the certainty of paternity (except in the case of sneaker males). Osteoglossum bicirrhosum is considered a monogamous fish, and has particular behavioral traits that permit the study of mating systems and parental care, such as male mouthbrooding. We investigated the genetic relationships of males with the broods found in their oral cavities in Osteoglossum samples collected in a natural environment in the lower Purus river basin, Amazonas, Brazil. Fourteen broods were analyzed for parentage (268 young and 14 adult males) using eight microsatellite loci. The results indicate that eleven broods show a monogamous system. In one brood, however, approximately 50% of the young were genetically compatible with being offspring of another male, and in another two broods, none of the subsampled young were compatible with the genotypes of the brooding male. The result of this first brood may be explained by the extra-parental contribution of a sneaker male, whereas cooperative parental care may explain the result in the other two broods.
... The result of this comparative study does not invalidate the present function of courtship feeding as paternal investment in bushcrickets. Males with large investments should have a high confidence of paternity (Werren et al. 1980;Gwynne 1984b), which assumes that the donating males will father the benefitted offspring (Gwynne 1988b). A paternal investment function may have arisen secondarily, once paternity assurance is high. ...
Article
During courtship and copulation, males of many insect species provide the female with a nuptial gift of a prey item or synthesized material. These gifts may be explained as a form of paternal investment by increasing female reproductive output, or in terms of mating effort by increasing male fertilization success. These explanations, while not mutually exclusive, are controversial. While experimental studies examine the maintenance of nuptial gifts in single species, comparative studies are required to indicate more general evolutionary trends. Male bushcrickets provide females with a nuptial gift, a spermatophylax, which is transferred to females at mating along with the sperm-containing ampulla. Analysis of comparative data of 28 species of bushcrickets (Orthoptera: Tettigoniidae), reveals that male spermatophore size (spermatophylax and ampulla weight) is positively correlated with female refractory period, which, in turn, correlates with male fertilization success. Moreover, gift size (the spermatophylax) covaries with ejaculate size (the ampulla), which is consistent with the hypothesis that it serves as a sperm protection device. In contrast, there is no significant correlation between any measure of female fecundity and male spermatophylax size. This indicates that the variation in spermatophore size among bushcrickets is better explained by a mating-effort function than a paternal investment function.
... In many socially monogamous and cooperatively breeding species, males' mating success may be constrained by such factors as the need for bi-parental care and female–female competition. Under such conditions, it may pay males to invest in current offspring, even if this comes at some expense to future mating opportunities (Maynard Smith 1977; Queller 1997; Werren et al. 1980). Our analysis suggests that this may also be true in some polygynous species. ...
Article
Full-text available
Sexual selection theory predicts that males in polygynous species of mammals will invest more reproductive effort in mate competition than parental investment. A corollary to this prediction is that males will mount a stress response when their access to mates is threatened. Indeed, numerous studies have shown that males exhibit elevated stress hormones, or glucocorticoids (GCs), when their access to females, or a proxy to this access like dominance rank, is challenged. In contrast, the relationship between stress hormones and paternal effort is less obvious. We report results from a study of wild male chacma baboons indicating that males experienced elevated GC levels during periods of social instability following the immigration of a dominant male. These effects were strongest in males whose mating opportunities were at greatest risk: high-ranking males and males engaged in sexual consortships. Males involved in friendships with lactating females, a form of paternal investment, also experienced high GC levels during these periods of instability. There was a tendency for males with lactating female friends to reduce their time spent in consortships during unstable periods, when the risk of infanticide was high. Thus, even in a highly polygynous mammal, males may have to balance paternal effort with mating effort. Males who invest entirely in mating effort risk losing the infants they have sired to infanticide. Males who invest in paternal care may enhance their offspring’s survival, but at the cost of elevated GC levels, the risk of injury, and the loss of mating opportunities.
... Electrical stimulation of the preoptic area in male bluegill sunfish also elicits paternal care (75). These results suggest that highly conserved circuits and neuroendocrine mechanisms may be repeatedly recruited to mediate similar social behaviors (2,76). What specific information is carried by these brain regions and how they encode the various components of parental care remain to be determined. ...
Article
Full-text available
Parental care, including feeding and protection of young, is essential for the survival as well as mental and physical well-being of the offspring. A large variety of parental behaviors has been described across species and sexes, raising fascinating questions about how animals identify the young and how brain circuits drive and modulate parental displays in males and females. Recent studies have begun to uncover a striking antagonistic interplay between brain systems underlying parental care and infant-directed aggression in both males and females, as well as a large range of intrinsic and environmentally driven neural modulation and plasticity. Improved understanding of the neural control of parental interactions in animals should provide novel insights into the complex issue of human parental care in both health and disease.
Article
Full-text available
Maternal effects are prevalent in nature and significantly contribute to variation in phenotypic trait expression. However, little attention has been paid to the factors shaping variation in the traits mediating these effects (maternal effectors). Specific maternal effectors are often not identified, and typically they are assumed to be inherited in an additive genetic and autosomal manner. Given that these effectors can cause long‐lasting effects on offspring phenotype, it is likely that they may also affect themselves in the next generation. Although the existence of such cascading maternal effects has been discussed and modeled, empirical examples of such effects are rare, let alone quantitative estimates of their strength and evolutionary consequences. Here, we demonstrate that the investment a mother makes in her eggs positively affects the egg investment of her daughters. Through reciprocally crossing artificially selected lines for divergent prenatal maternal investment in Japanese quail (Coturnix japonica), we demonstrate that the size of eggs daughters lay resembles the egg size of their maternal line significantly more than that of their paternal line, highlighting that egg size is in part maternally inherited. Correspondingly, we find that variation in the daughters' egg size is in part determined by maternal identity, in addition to substantial additive genetic effects. Furthermore, this maternal variance in offspring egg size is fully explained by maternal egg size, demonstrating the presence of a positive cascading effect of maternal egg size on offspring egg size. Finally, we use an evolutionary model to quantify the consequences of covariance between cascading maternal and additive genetic effects for both maternal effector and offspring body mass evolution. Our study demonstrates that by amplifying the amount of variation available for selection to act on, positive cascading maternal effects can significantly enhance the evolutionary potential of maternal effectors and the offspring traits that they affect.
Chapter
This chapter focuses on a related but less widely recognized phenomenon, indirect genetic effects (IGEs), that also blurs the traditional distinction between genetic and environmental effects. It focuses primarily on IGEs affecting the expression of behavioural traits and provides a discussion on IGEs arising from interactions among unrelated individuals. The chapter provides a brief introduction to traditional quantitative genetic theory. It shows how the quantitative genetic theory can be extended to include social interactions and illustrates what consequences this has for the evolution of behaviour. IGEs represent an additional source of genetic variance for behaviours that change our expectations, relative to those derived from direct genetic effect (DGE)‐only models, of phenotypic responses to selection. A common outcome of most theoretical models is that IGEs will influence the evolutionary dynamics of phenotypic traits involved in the social interactions.
Article
Full-text available
Parent and offspring behaviours are expected to act as both the agents and targets of selection. This may generate parent‐offspring coadaptation in which parent and offspring behaviours become genetically correlated in a way that increases inclusive fitness. Cross‐fostering has been used to study parent‐offspring coadaptation, with the prediction that offspring raised by non‐relatives, or parents raising non‐relatives, should suffer fitness costs. Using long‐term data from more than 400 partially crossed broods of blue tits (Cyanistes caeruleus) we show there is no difference in mass or survival between crossed and non‐crossed chicks. However, previous studies for which the evidence for parent‐offspring coadaptation is strongest compare chicks from fully crossed broods with those from non‐crossed broods. When parent‐offspring coadaptation acts at the level of the brood then partial cross‐fostering experiments are not expected to show evidence of coadaptation. To test this, we performed an additional experiment (163 broods) in which clutches were either fully crossed, non‐crossed, or partially crossed. In agreement with the long‐term data, there was no evidence for parent‐offspring coadaptation on offspring fitness despite high power. In addition there was no evidence of effects on parental fitness, nor evidence of sibling coadaptation, although the power of these tests was more modest. This article is protected by copyright. All rights reserved
Article
To infer that a single‐nucleotide polymorphism (SNP) either affects a phenotype or is linkage disequilibrium with a causal site, we must have some assurance that any SNP‐phenotype correlation is not the result of confounding with environmental variables that also affect the trait. In this study, we study the properties of linkage disequilibrium (LD) Score regression, a recently developed method for using summary statistics from genome‐wide association studies to ensure that confounding does not inflate the number of false positives. We do not treat the effects of genetic variation as a random variable and thus are able to obtain results about the unbiasedness of this method. We demonstrate that LD Score regression can produce estimates of confounding at null SNPs that are unbiased or conservative under fairly general conditions. This robustness holds in the case of the parent genotype affecting the offspring phenotype through some environmental mechanism, despite the resulting correlation over SNPs between LD Scores and the degree of confounding. Additionally, we demonstrate that LD Score regression can produce reasonably robust estimates of the genetic correlation, even when its estimates of the genetic covariance and the two univariate heritabilities are substantially biased.
Article
Full-text available
Co-operative breeders provide parental care to non-filial offspring—a behaviour known as ‘alloparental care’. While inclusive fitness benefits are a widely accepted driver of alloparental care in kin-based social groups, such indirect benefits are lost in non-kin societies. Among such societies, theory predicts that the degree of parental and alloparental effort should therefore be proportional to an individual’s genetic contribution to mixed broods—depending upon reproductive options. Using genotyping data across five to 12 microsatellite loci for individuals from 20 social groups (67 adults and 153 nestlings), we assessed whether kinship or proportional reproductive success explained trends in parental and alloparental effort in the Smooth-billed Ani (Crotophaga ani), a joint-nesting cuckoo species. Nocturnal incubation in this species appears to be performed almost exclusively by a single male. We first report significantly higher degrees of relatedness between adults within social groups (\(\bar{r}\) = 0.208, n = 114 dyads), than between social groups (\(\bar{r}\) = 0.120, n = 893 dyads), suggesting that inclusive fitness benefits may in part explain uneven allocation of parental effort. Second, we show that nocturnal incubation status is a significant predictor of reproductive success in males, as nocturnal incubators sire a greater proportion of nestlings in mixed-parentage broods. While patterns of reproductive skew appear high at 80% paternal confidence (\(\bar{B}\) = 0.052, p = 0.061), we report no significant deviation from an egalitarian breeding framework. Our results revealed similar patterns of reproductive allocation to closely related Groove-billed Anis (Crotophaga sulcirostris); however, differences in male reproductive skew and within-group relatedness across crotophagids are highlighted and offer insight into social evolution among anis.
Article
The pattern of sperm predominance in doubly mated female crickets, Gryllodes supplicans, was investigated using a radiation-sterility technique. Female G. supplicans made significant use of sperm from both males in fertilizing eggs; overall, first males to mate enjoyed a small advantage, fertilizing about 60% of the offspring produced subsequent to the second mating. The combined use of the sperm of both males in fertilizing eggs occurred soon after the second mating; evidently, mixing of ejaculates within a female's spermatheca does occur. Male G. supplicans provide females with a nuptial gift, the spermatophylax, which influences the time at which a female removes the externally attached sperm-ampulla; this in turn determines the quantity of sperm that is transferred. Moreover, the degree of sperm precedence achieved by a male may be positively related to the time at which the female removes his sperm ampulla. Thus males, by feeding females, ensure not only that a sufficient number of sperm are transferred to fertilize all of a female's eggs, but also may increase the certainty of their paternity. In mating systems in which females control sperm transfer and paternity is influenced by numbers of sperm (i.e., numerical sperm competition), an increase in prezygotic investment in females may be an adaptive male response.
Article
There is abundant evidence in many taxa for positive directional selection on body size, and yet little evidence for microevolutionary change. In many species, variation in body size is partly determined by the actions of parents, so a proposed explanation for stasis is the presence of a negative genetic correlation between direct and parental effects. Consequently, selecting genes for increased body size would result in a correlated decline in parental effects, reducing body size in the following generation. We show that these arguments implicitly assume that parental care is cost free, and that including a cost alters the predicted genetic architectures needed to explain stasis. Using a large cross-fostered population of blue tits, we estimate direct selection on parental effects for body mass, and show it is negative. Negative selection is consistent with a cost to parental care, mainly acting through a reduction in current fecundity rather than survival. Under these conditions, evolutionary stasis is possible for moderately negative genetic correlations between direct and parental effects. This is in contrast to the implausibly extreme correlations needed when care is assumed to be cost free. Thus, we highlight the importance of accounting correctly for complete selection acting on traits across generations. This article is protected by copyright. All rights reserved.
Chapter
During the past few years interest among primatologists in male parental care has increased. In this paper I will describe a series of events which occurred in the Arashiyama B troop of Japanese macaques in the spring of 1978 that resulted in the third highest ranking male, Ran63, becoming the primary caretaker of two sisters— one 10 months of age and the other 22 months of age at the time of the adoption — and another unrelated male infant also 10 months of age at the time of the adoption. The term primary caretaker is being used here to define a situation in which (1) the mother is no longer available to be the caretaker, (2) the adoptee(s) maintain close proximity to the male while traveling, foraging or eating provisioned food and may even be carried by the male (although Ran63 did not in this case carry any of his adoptees), (3) the male protects the adoptee(s) from the aggression of other troop members, and (4) the male and the adoptee(s engage in grooming and huddling while resting and sleeping. Following the description of the adoptions, a series of questions involving altruism, kin selection, and reproductive strategies will be explored. It is hoped that by exploring these questions the behavior of Japanese macaque males toward motherless infants and juveniles will be better understood. Because of the rarity of such adoptions, the data are necessarily anecdotal. As an aside, adult Arachiyama females have not been observed adopting motherless infants.
Chapter
In the last several years the Nature and origins of paternal care have become increasingly important to many people in Western societies. Our own culture is currently undergoing sweeping challenges to the traditional sex roles maintained since the late 19th century. Women today are more frequently seeking employment outside the home, both for personal fulfillment and for economic reasons, and the role of fathers in caring for children has become more critical for many families. As we seek to alter our own traditions of paternal care, we may be able to gain some perspective from studies of paternal care in animals, especially paternal care in Primates, those animals closest to us phylogenetically.
Article
The chapter describes parental care in intertidal fishes by addressing two themes: if there is something unique about parental care in this group, and if intertidal fishes can help to understand the broader issues of parental care in fishes. Parental care is defined as any investment by the parent in an individual offspring that increases the offspring's chance of surviving at the cost of the parent's ability to invest in other offspring. In fishes, parental care can take a variety of forms, including guarding, nest building and maintenance, substrate cleaning, fanning, internal gestation, removal of dead eggs, oral brooding, retrieval of eggs or fry that stray from the nest or school, cleaning of eggs or fry, external egg carrying and splashing. Guarding is the predominant form of care, occurring in 95% of care-giving fishes. Uniparental care is most common (78% of families providing care have solitary male or solitary female care). Male care is more common than female parental care (61% of families showing care have male care).
Article
Full-text available
The parental behavior of Microtus socialis Pall, males and females depending on their preliminary experience in litter rearing was studied in laboratory. All the pairs cohabited permanently in one nest. The males exhibited all the patterns of direct care of pups except for nursing: huddling over, brooding with kyphosis, grooming, and other ones. There were no differences between different sexes in the total time spent in the nest, as well as in the time spent alone there. However, the females groomed the pups much longer that males did. The rate of manipulation with nest material was much higher in experienced females than in inexperienced ones, and experienced males groomed pups longer than inexperienced ones. The parental behavior of social voles was compared with that of other vole species. The importance of males contributed to the parental care is discussed, especially to the haptic stimulation of pups, as well as to the formation of stabile family-group mode of life among rodents is discussed.
Article
The pattern of sperm predominance in doubly mated female crickets, Gryllodes supplicans, was investigated using a radiation-sterility technique. Female G. supplicans made significant use of sperm from both males in fertilizing eggs; overall, first males to mate enjoyed a small advantage, fertilizing about 60% of the offspring produced subsequent to the second mating. The combined use of the sperm of both males in fertilizing eggs occurred soon after the second mating; evidently, mixing of ejaculates within a female's spermatheca does occur. Male G. supplicans provide females with a nuptial gift, the spermatophylax, which influences the time at which a female removes the externally attached sperm-ampulla; this in turn determines the quantity of sperm that is transferred. Moreover, the degree of sperm precedence achieved by a male may be positively related to the time at which the female removes his sperm ampulla. Thus males, by feeding females, ensure not only that a sufficient number of sperm are transferred to fertilize all of a female's eggs, but also may increase the certainty of their paternity. In mating systems in which females control sperm transfer and paternity is influenced by numbers of sperm (i.e., numerical sperm competition), an increase in prezygotic investment in females may be an adaptive male response.
Article
Some have noted that hunting by males may represent a strategy of "showing off" rather than a strategy of "maximizing efficiency." Others have observed that this insight is limited by insufficient theory for predicting when men should pursue one strategy instead of the other. This article attempts to foster development of such theory by more explicitly casting the issue in terms of the trade-off between mating effort and parenting effort and by discussing conditions under which males benefit by investing more in one area than the other. Testing predictions derived from this framework should help resolve a current debate among scholars of California and Great Basin prehistory.
Article
In cooperative breeding systems, males that share a nest face the prospect of providing for young that are not their own. Males of many species attempt to reduce the risk of losing paternity with aggressive behaviors, thereby limiting other males' access to the female during copulation. The Galapagos Hawk (Buteo galapagoensis) exhibits an extreme form of cooperative polyandry in which anecdotal data suggest all males in a territory share mating equally with the female, with very little to no interference, and care for young within the group. Males in a territory are unrelated adults and share paternity. We examine paternal care in relation to the shared parentage of the Galapagos Hawk and offer explanations for group cohesion. We found that paternal care was variable and that all males cared for the young on their territory without regard to the number of males residing together. There was evidence that males that sired young and those that sired none did not differ in quantity of care. However, we could not rule out a relationship between paternity and care. There was no obvious cue the males could use to discern paternity, since the only evidence of dominance was a subtle hierarchy expressed in larger groups. We suggest that the simple rule for paternal care in the Galapagos Hawk is that ifa male is a group member, he will copulate with the female, have some probability to fertilize the eggs, and provide care for young produced at the nest.
Article
Full-text available
The zebra cichlid Cichlasoma nigrofasciatum is a species famous for its behaviors of biparental care. This study was conducted to examine the effect of parental care and their size on the reproductive success of this fish. Hatching rate (HRs) has significant difference in four parental care treatment, 1. bi-parental, 2. female-only, 3. male-only and 4. no-parental care. Mean hatching rate was 94.81, 93.36, 86.15 and 84.44, respectively. There was no significant difference in HRs between female-only and bi-parental care as well as no-parental with male-only care. Also, there were significant positive correlations of egg numbers (ENs) but not HRs to standard lengths (SLs) of parent. These results suggest that reproductive success of this fish primarily depends on bi-parental care and greater size.
Article
Full-text available
In light of current knowledge of the behavioral and genetic complexity of avian mating systems, both conventional classifications of mating systems and traditional models for their evolution are inadequate. In this paper we attempt to provide a synthetic view of the major avian mating systems, recognizing common aspects without imposing arbitrarily rigid categories. Our scheme incorporates both genetic and social aspects of mating systems and focuses on individual mating tactics. Although classification tends to imply stasis, we suggest that mating systems may be quite fluid in both ecological and evolutionary time. Flow is possible because the range of mating tactics available to both male and female birds is not limited by the mating system. Thus, individuals may respond quickly to the opportunities and constraints of changing circumstances. The movement of populations or species between mating system types may also be closely related to the fluctuating heritability of traits used in mate choice. We suggest that conditional mating tactics and female mating (as opposed to parental) tactics are key mating system parameters. Predictions (such as the expected relationship between paternity and male parental investment) that ignore conditional mating tactics, or explanations (such as for consortship patterns) that ignore conditional or female mating tactics are likely to be simplistic or erroneous. Future understanding of avian mating systems will be most enhanced by close analysis of the opportunities and activities of all phenotypic classes within a population, and by experimental manipulation of those opportunities. Genetic, developmental, and historical constraints that may affect the expression of mating tactics should also receive more attention. As this kind of information becomes available, a more complete, accurate, and integrated understanding of avian mating systems will be possible.
Article
Experimental crosses with and without risk of sperm competition were performed in the locust Locusta migratoria, an orthopteran species with a certain degree of first male sperm precedence. The results showed that first mating males perform a significantly higher number of matings than non-first ones, which is paralleled to a shorter remating period. Possible explanations of this differential male mating frequency are discussed in the light of current hypotheses on sexual selection theory.
Article
In polygynous species, males appear to gain additional offspring by pairing with multiple females simultaneously. However, this may not be true if some females copulate outside of the social pair bond. Polygynous males could experience lower paternity because of trade-offs among gaining multiple social mates, guarding fertility with these mates, and pursuing extra-pair matings. Alternatively, polygynous males could simultaneously gain extra social mates and have high paternity, either because of female preferences or because of male competitive attributes. We tested four predictions stemming from these hypotheses in a facultatively polygynous songbird, the dickcissel (Spiza americana). Unlike most previous studies, we found that males with higher social mating success (harem size) also tended to have higher within-pair paternity and that the number of extra-pair young a male sired increased significantly with his social mating success. Females that paired with mated males were not more likely to produce extra-pair young. In contrast, extra-pair paternity was significantly lower in the nests of females whose nesting activity overlapped that of another female on the same territory. This pattern of mating was robust to differences in breeding density. Indeed, breeding density had no effect on either extra-pair mating or on the association between polygyny and paternity. Finally, nest survival increased with harem size. This result, combined with the positive association between polygyny and paternity, contributed to significantly higher realized reproductive success by polygynous male dickcissels.
Article
Males of the bushcricket, Requena verticalis, contribute parental investment at mating via a nutrient-rich spermatophore. Because the first male to mate has a high confidence of paternity, subsequent males have their investment cuckolded. The effects of first male sperm precedence on male mate choice and protandry were investigated. Males were equally likely to mate with virgin and non-virgin females during mating trials. However, they could discriminate on the basis of female age and mated preferentially with young females in laboratory trials and in the field. By using age as a cue, males increase the probability of rejecting non-virgin females, thereby reducing the risks of cuckoldry. It is argued that failure to recognize non-virgins per se may have arisen by sexual conflict over mating, and that the high degree of protandry is an adaptive strategy by males to mate with their preferred young females and enhance their confidence of paternity.
Article
Nest attendance by male and female meadow voles, Microtus pennsylvanicus , was monitored in 24-h recordings. Fathers housed with mothers and their pups, but with no other adults present, were in the natal nest for much of the observation period. When unfamiliar males were present, instead of, or in addition to, the fathers, females prevented the unfamiliar males from entering the natal nest, and showed reduced tolerance to the fathers' attempts at nest entry. Females housed with unfamiliar males were the only voles to spend more time in the nest at night than during the day, which may be related to the only instance of infanticide occurring when an unfamiliar male successfully entered the nest at night. The introduction of an oestrous female did not reduce the amount of time the father spent in the nest, although several fathers mated with the oestrous females. In another experiment, females were found to be temporarily less tolerant of fathers that were removed during pregnancy and returned after the pups were born, but after renewed contact, the females allowed the males to enter the nest. Fathers tested in the absence of mothers spent a substantial portion of the test time with pups, and effectively prevented unfamiliar males from gaining access to their pups. These results were related to the seasonal changes in nest sharing in meadow voles, suggesting that male reproductive tactics should be flexible to accommodate changes in mating opportunities and changes in the value of biparental care.
Article
Based on the model of Whittingham et al. (1992, Am. Nat., 139, 1115-1125), it is predicted that males in many monogamous species of birds will not decrease their level of parental care until their confidence of paternity is very low (threshold response). This prediction was tested by holding monogamous male tree swallows, Tachycineta bicolor, in captivity for 1 or 3 days during their mate's fertile (pre-laying, laying) or non-fertile (incubation) periods. Males were held in cages with one-way glass that allowed them to see their mates engage in extra-pair copulations. Females increased their rate of extra-pair copulation when their mates where held in captivity during pre-laying and laying. Therefore, the confidence of paternity of these captive males was lower than unmanipulated males or males held during incubation (controls). Although the male's access to his fertile mate was reduced by up to 57%, there was no effect of the timing or length of captivity on the male's share of feeding nestlings or his defence of the nest. Male tree swallows did not reduce their parental effort over a wide range of paternity, and previous studies showed they commit infanticide when their paternity is zero, suggesting that they exhibit a threshold response to decreasing paternity. Furthermore, a review of several studies suggests that other monogamous species also exhibit a threshold response to decreasing paternity.
Article
Parentage is the proportion of juveniles in a brood that are offspring of potential care givers. We analyzed how reductions in parentage affect the evolution of parental behavior using a static optimization model. The main benefit of parental effort was an increase in the survival of offspring, and the main costs were reduced opportunities to seek additional matings or to parasitize neighbors and/or reduced survival. Both the costs and benefits included terms for relatedness to young. The effect of parentage depended on (1) whether parents responded in ecological time (facultative response) or in evolutionary time (nonfacultative response), (2) whether the cues enabling assessment of parentage permitted discrimination among off- spring, and (3) whether parentage was the same among different groups of juveniles (unrestricted) or varied between them (restricted). When parents did not know their own parentage and mean parentage was the same for all matings, reduced parentage affected the costs and benefits equally, so, as in several previous models, there was no effect on the optimal level of parental effort. Parentage did affect optimal parental effort when mean parentage to the present brood differed from that to young from alternative or future matings. Lowered parentage reduced optimal parental effort when the cost of parenting was missed opportunities for extrapair copulations or brood parasitism or when parentage was consistently higher in alternative or future matings. Nonlinear changes in parentage with age gave complex trajectories of parental care, with individuals of different ages having similar parentage but exhibiting different levels of parental effort. Correlations between parentage and other variables in the model (such as opportunities for additional matings) sometimes masked, but never eliminated, the effects of parentage. When parents could discriminate their own young in a brood, overall parental effort was reduced, but nepotism was increased. When parents could not discriminate their own offspring but had general cues about average parentage to the brood, effects varied depending on the costs and benefits of parental behavior. When parental behavior was costly to care givers, parentage had more effect than when parenting was not costly. Likewise, parentage had less effect when care greatly increased offspring survival than when care was less necessary. Our analyses reconcile conflicting results from previous models and suggest a general framework for analyzing parental behavior within populations and among higher taxonomic groups. Key words: paternity, maternity, parental investment, uncertain parentage, relatedness, optimization models, extrapair copulations, intraspecific brood parasitism. (Behav Ecol 4:66-77 (1993))
Article
The phenomenon of male parental care, which is unusually common in bony fishes, has been the subject of numerous evolutionary hypotheses in recent years. In an effort to evaluate these hypotheses the results of a survey of all families whose species exhibit parental care is presented. Sexual selection and female choice for male parental behavior only partially explain this phenomenon. Although females may benefit by males being the care-giving individuals, these benefits may apply equally to males when females are the care-givers. The information avaliable suggests that the effects of care-giving on the future reproduction of the male and the male's provability of genetic relatedness to his mate's offspring are major factors in the evolution and maintanance of male parental behavior. The costs of care-giving are minimized by mating strategies thath enagle males to give parental care and pursue further mating simulteneously. Male fishes with external fertilization often attain a relatively high probability...
Article
Sexual roles in brood care have been examined for a number of animal groups. Most authors have related these patterns to general ecological characteristics of the animal's environment, such as distribution and character of food, nature of predation, and difficulty of finding and keeping a mate (e.g., Verner and Willson 1966, for birds; Orians 1969, for birds and mammals; Crook 1970, for primates; Kleiman and Eisenberg 1973, for canids and felids; Alexander 1974, for social animals). A second approach has been that of Trivers (1972) and Williams (1975), who explored the relation between parental care patterns and mating systems. Trivers pointed out that reproductive effort is divided between courtship (i.e., efforts to get mates) and parental investment, (PI), that is, efforts to improve the probability of an individual offspring's survival. The latter limits the parent's ability to invest in other offspring, current and future. Trivers also suggested that maximum reproductive success would often result in different courtship and parental roles for the sexes, and that this provided the most basic explanation for the variety of mating patterns found in nature. Williams reached similar conclusions, and provided examples from a variety of taxonomic groups illustrating differences between the sexes in the form of parental investment. The roles of the sexes in brood care in fishes differ among species, and although some authors have considered them peripherally, at present no comprehensive theory has been developed to explain them. In this paper we develop a theory to account for the sexual roles in different fish parental care patterns based on the approaches of Trivers (1972) and Williams (1975).
Article
Usually working in pairs, Nicrophorous beetles rapidly bury carcasses weighing up to 100 gms. Their behavior shows considerable plasticity. The beetles can transport a carcass to a suitable burial site and are able to overcome a variety of obstacles. The buried carcasses provide food for the beetles and their larvae. Patterns of social cooperation are described. (PsycINFO Database Record (c) 2012 APA, all rights reserved)
Article
I . Intrasexual seiection may have played a large part in insects in the evolution of copulation with internal fertilization from indirect spermatophore-transferring acts. 2. ‘Sperm competition’ may be defined as the competition within a single female between the sperm from two or more males over the fertilization of the ova. 3. There is considerable evidence from sperm-marking experiments that sperm competition is very common in insects as a result of multiple matings. Insects so far examined show that sperm from all inseminations can be used (to a varying extent) in the fertilization of subsequent offspring, but mating does not always result in successful insemination. In most cases (so far examined), the last male to mate tends to predominate in fertilizing the offspring. 4.Insects are preadapted to sustaining a very high level of sperm competition, compared with several other animal groups. The main preadaptations may be sum- marized as follows: (a)Females often mate several times within the duration of effec- tiveness of a given ejaculate. There may be several reasons for this. Though most usually females are unreceptive for some time after mating, some species appear ‘promiscuous’. Unreceptive females are also sometimes raped. Male persistence is sometimes prolonged, so that full female receptivity is advantageous. It is advan- tageous for a female to become receptive again when fertility first begins to decline; this is not when all the first ejaculate is used up. (b) Female insects typically possess specialized sperm storage organs in which sperm can be maintained in a viable condi- tion for a very long time, often until the death of the female. (c) Extremely efficient utilization of stored sperm at the time of fertilization appears to be an insect charac- teristic. In Drosophila the number of stored sperm virtually equals the possible number of fertilizations. Overlapping of effective ejaculates is therefore high. Second insemina- tions almost invariably reduce the fertility which would have been experienced by the first male to mate. 5. It is argued that this preadaptation to a very high level of sperm competition has led to intense intrasexual selective pressures on the male. In response to these pressures, two main lines of sexually selected adaptation are predicted: (a) towards mechanisms by which a male inseminates a female in such a way as to achieve pre- cedence over previously stored sperm and (b) accentuated by the above adaptation, mechanisms will evolve by which a male which mates with a given female will reduce the occurrence or success of subsequent inseminations with that female. These two forms of adaptation are diametrically opposed ; a high selective advantage would be gained by a male which superseded previous sperm and prevented any subsequent successful inseminations. 6. Several adaptations in male insects can be interpreted in the light of the above predictions, though many of these may also have other adaptive values through natural selection. The adaptation which evolves is not necessarily that which yields the maxi- mum possible egg gain to a given male (i.e. total sperm precedence), but that which results in the highest fertilization rate. 7. Sperm precedence is achieved in Drosophila by sperm displacement, where sperm from a second male predominate over previously stored sperm by directly displacing them from the sperm stores. Sperm displacement may occur in many insects. 8. Several behavioural and physiological adaptations of male insects may help to reduce the effectiveness, or occurrence of second inseminations of the same female by other males. These include : ( a ) Mating plugs (sphragis, spermat0phragmata)-male accessory gland secretions, usually transferred after insemination, which coagulate and form plugs within the female genital tract. Plugs may often serve to ‘guard’ the female until unreceptivity is initiated. In many insects, agents in the seminal fluid or male accessory gland secretions induce unreceptivity in the female. (b)Prolonged copu- lation-sometimes copulation takes much longer than seems necessary merely to transfer the sperm. This may have the same functions as that of a mating plug, but renders the male unfree to search for further females. (c) Passive phases (amplexus, tandem behaviour)-stages of the male’s reproductive behaviour during which he remains mounted on or otherwise attached to the female but without true genital contact between the two sexes. Postcopulatory passive phases sometimes serve to guard the female during oviposition where high densities of searching males are pre- valent. The postcopulatory passive phase of Scatophaga has an extremely high intra- sexual selective advantage which exceeds any apparent natural selective advantage by two orders of magnitude. (d)Non-contact guarding phases-reproductive behaviour phases during which the male remains close but not in contact with the female, guard- ing her from other males. Postcopulatory non-contact guarding phases appear to have the same selective advantage as postcopulatory passive phases. 9. Mechanisms to avoid ‘take-over ’ during copulation, passive, and non-contact guarding phases also serve to reduce sperm competition. These include increased efficiency of grasping apparatus, specialized rejection reactions which serve to dispel or ‘trick’ the recognition mechanism of the attacker, and emigrations from the site of highest probability of ‘take-over ’. There is quantitative evidence in Scatophaga that the emigration threshold of copulating males is determined by this form of intrasexual selective pressure. 10.Precopulatory passive phases may serve mainly to keep the sexes together until the female becomes receptive, but share several features in common with postcopula- tory passive phases. Territoriality of male insects may have arisen primarily through sexual selection as a mechanism by which a male guards an area into which a female is most likely to enter.
Article
Estimates of body weight, group size, home range size, day range length, socionomic sex ratio and sexual dimorphism are compared between 100 primate species, allocated to seven ecological categories. As would be predicted on energetic grounds, home range size and day range length are positively related to group weight and are greater in frugivores than in folivores; population density is negatively related to body weight; and group size is positively related to body weight. The adaptive significance of Variation in body size, sexual dimorphism and socionomic sex ratio is also discussed.
Article
TRIVERS1 defines parental investment as ``any investment by the parent in an individual offspring that increases the offspring's chance of surviving ... at the cost of the parent's ability to invest in other offspring''. He has cleverly used this definition to elucidate many outstanding problems in social ethology1-3. But one of his most appealing usages of it is fallacious.
Article
The science of sociobiology, which began in principle with the work of Fisher and Haldane and has more recently been developed by Hamilton, Maynard Smith, Trivers, Wilson and others, has been the centre of both scientific and political controversy. Dr Parker discusses the strengths and weaknesses of the approach, and illustrates that behaviour can be adapted in a complex way in conformity with sociobiological theory.
Article
This review considers the behavioral, ecological, and reproductive characteristics of mammals exhibiting monogamy, i.e., mating exclusivity. From a discussion of the life histories of selected species of monogamous primates, carnivores, rodents and ungulates, several trends emerge. Two forms of monogamy occur, Type I, facultative, and Type II, obligate. The selective pressures leading to these two forms of monogamy may have been different. Facultative monogamy may result when a species exists at very low densities, with males and females being so spaced that only a single member of the opposite sex is available for mating. Obligate monogamy appears to occur when a solitary female cannot rear a litter without aid from conspecifics, but the carrying capacity of the habitat is insufficient to allow more than one female to breed simultaneously within the same home range. Within both types of monogamy, the following traits are typically seen: (1) adults show little sexual dimorphism either physically or behaviorally: (2) the adult male and female exhibit infrequent socio-sexual interactions except during the early stages of pair bond formation. Additional trends specific to mammals exhibiting obligate monogamy are: (1) the young exhibit delayed sexual maturation in the presence of the parents, and thus only the adult pair breeds; (2) the older juveniles aid in rearing young siblings; and (3) the adult male (father) aids in the rearing of young by any or all of the following: carrying, feeding, defending, and socializing offspring.
Article
A genetical mathematical model is described which allows for interactions between relatives on one another's fitness. Making use of Wright's Coefficient of Relationship as the measure of the proportion of replica genes in a relative, a quantity is found which incorporates the maximizing property of Darwinian fitness. This quantity is named “inclusive fitness”. Species following the model should tend to evolve behaviour such that each organism appears to be attempting to maximize its inclusive fitness. This implies a limited restraint on selfish competitive behaviour and possibility of limited self-sacrifices. Special cases of the model are used to show (a) that selection in the social situations newly covered tends to be slower than classical selection, (b) how in populations of rather non-dispersive organisms the model may apply to genes affecting dispersion, and (c) how it may apply approximately to competition between relatives, for example, within sibships. Some artificialities of the model are discussed.
Article
Male ring doves exhibit less courtship and more aggressive behavior toward females that have recently associated with other males than to females that have been isolated. The difference in response may be related to the differing probability of cuckoldry.