Large males have a mating advantage in a species of darter with smaller, allopaternal males Etheostoma olmstedi

Theory suggests that males that are larger than their competitors may have increased mating success, due to both greater competitive ability and increased attractiveness to females. We examined how male mating success varies with male size in the tessellated darter Etheostoma olmstedi. Previous work has shown that large males tend to move around and breed in vacant breeding sites, and consequently provide less care for their eggs, while smaller individuals can be allopaternal, caring for the eggs of other males as well as for their own. We studied female egg deposition in a natural breeding population using artificial breeding sites and in the laboratory, where female choice of spawning site was restricted to two breeding sites tended by two males of different sizes. In both the field and the laboratory, nests tended by larger males were more likely to receive new eggs. Additionally, the mean size of males associated with a nest was positively correlated with both the maximum coverage of eggs at the nest and the number of times new eggs were deposited. We discuss how the increased mating success of larger males, despite their decreased parental care, may help explain allopaternal care in this species [Current Zoology 56 (1): 1-5, 2010].     

Short‐term dynamics of nest occupancy in an allopaternal species,the tessellated darter Etheostoma olmstedi

To inquire how male size interacts with alloparental behaviour and mating success in the tessellated darter Etheostoma olmstedi, males were given a choice of nests with or without eggs; subsequent nest occupancy, takeovers and egg deposits were monitored. Subordinate males readily occupied available nests with eggs but were often evicted by dominant males, suggesting that males of all sizes compete for the opportunity to provide allopaternal care in this species.     

Allopaternal care in the tessellated darter,Etheostoma olmstedi (Pisces: Percidae)

Synopsis Large male tessellated darters, Etheostoma olmstedi, defend flat-bottomed rocks, the undersides of which serve as spawning substrates. Because females attach eggs directly onto bare stone, a spawning bout necessarily decreases the surface area available for further breeding at a nest. In apparent response to the female preference of ovipositing on rocks with the most bare surface, fathers abandon nests with their attached offspring to search for rocks with more uncovered surface. Egg abandonment also results after roving fathers intercept and spawn with ripe females and remain at a different rock to care for a new clutch. In essence, large males appear to maximize the number of eggs they fertilize by sequentially monopolizing breeding patches that are temporarily most desirable to females. The costs to a father of abandoning his eggs appear to be low for two reasons: (1) non-territorial males, created by a scarcity of suitable rocks, readily occupy vacant nests, and clean and aerate the abandoned eggs; and (2) after water hardening, eggs are less vulnerable to cannibalism. The maintenance of eggs by nonparental males may be both incidental to scrubbing the ceiling for their own breeding and adaptive in that viable eggs stimulate females to spawn; neither would appear to involve parental investment costs. Thus, allopaternal care in the tessellated darter may have evolved because it is performed in the selfish pursuit of spawning opportunities while entailing little, if any, of the costs normally subsumed within parental investment.     

Nest defense and aggressive interactions between a small benthic fish (the johnny darter Etheostoma nigrum) and crayfish

Synopsis Aggression by nest-guarding male johnny darters, Etheostoma nigrum, against intruding crayfish was investigated in laboratoy experiments and field observations. In the laboratory, darter success in chasing crayfish, Orconectes rusticus, from the nest site was inversely related to crayfish size. Small crayfish (less than 15 mm carapace length) were routinely evicted from the nest area by nips directed at the posterior end of the abdomen. Although such aggressive behavior was less successful against larger crayfish, even the largest crayfish tested (carapace length 30–32 mm) were chased from the nest area in 33% of the trials. Those large crayfish that entered nests often remained despite repeated attacks by the male johnny darter and egg predation was observed. In a small Ohio stream, openings to johnny darter nests were generally between 7 and 13 mm. Thus crayfish with a carapace height greater than 13 mm (corresponding to a carapace length greater than 29 mm) would not be able to enter johnny darter nests. In field observations, male Johnny darters successfully defended nests against another crayfish species (Orconectes sanborni, carapace length 12–29 mm). Together, aggressive behavior and small size of nest entrances allow the johnny darter to successfully reproduce in areas with abundant crayfish. The Unit is sponsored jointly by the United States Fish and Wildlife Service, The Ohio Department of NaturalResources, and The Ohio State University.     

Temporal and spatial patterns of spawning site use by two cavity-spawning darters in a Wisconsin stream

Synopsis There was partial spatial and complete temporal overlap of spawning by Etheostoma flabellare and E. nigrum in Marsh Creek, Wisconsin. E. flabellare tended to spawn at shallower depths than E. nigrum. Both species spawned over a greater range of depths in an area with few spawning sites (cavities under rocks or logs) than in an area where spawning sites were abundant. In the laboratory E. flabellare were able to prevent E. nigrum from occupying cavities they would have otherwise occupied. E. nigrum appear to be competitively inferior to E. flabellare in terms of spawning-site acquisition and retention, but possess several attributes that may allow them to persist in the presence of the latter.     

Filial cannibalism in a non-brood cycling marine fish

Synopsis Direct observation and the analysis of stomach contents indicate that custodial male Cortez damselfish, Stegastes rectifraenum, regularly eat some of the eggs they fertilize and guard. The pattern of forced starvation and brood cycling by custodial males that is seen in sticklebacks and other fishes for which filial cannibalism is known in the field does not exist for this species. These results indicate that filial cannibalism is likely to be more common among fishes than previously thought, because this behavior is not restricted to species that brood cycle.     

Infant development and parental care in two species of sifakas

This paper reports the results of a three-month field study on parental care and infant development in the diademed sifaka (Propithecus diadema edwardsi) in the primary rain-forest of south-eastern Madagascar. They are compared with a three-and-a-half-month study of captive white sifaka (Propithecus verreauxi coquereli) in the Duke University Primate Centre. Records were taken by means of focal animal and instantaneous sampling. In both species the mother was the primary carrier and caretaker; theP. verreauxi father carried the infant significantly more than did any animal other than the mother in theP. diadema group. The infantP. verreauxi spent less time off the mother than didP. diadema from week 4 through week 10. It is concluded thatP. verreauxi shows more non-maternal care thanP. diadema and also develops at a slower rate. The difference in the two species' habitats is discussed as a possible cause.     

Lack of sexual experience does not reduce the responses of LH, estrus or fertility in anestrous goats exposed to sexually active males

We investigated whether LH secretion, estrous behavior and fertility would differ between sexually inexperienced and experienced anestrous goats exposed to the males. Male goats were rendered sexually active during the reproductive rest season by exposure to 2.5 months of artificial long days. Two groups of anovulatory sexually inexperienced and sexually experienced does were exposed to males during 15 days (n = 20 per group). LH pulsatility was determined every 15 min from 4 h before to 8 h after introducing males (Day 0). Estrous behavior was recorded twice daily. Pregnancy rates were determined on Day 50. Fertility was determined at parturition. Male sexual behavior was registered on days 1 and 2 during 1 h. Before introducing the males, the number of LH pulses did not differ between groups. After introduction of the males, all females increased their LH pulsatility, but the number of pulses did not differ between sexually inexperienced and experienced goats. The proportion of females displaying estrous behavior with a high pregnancy rate and fertility did not differ between inexperienced and experienced goats. The sexual behavior of the males did not differ significantly between those interacting with sexually inexperienced or experienced goats. We conclude that goats can show substantial endocrine and reproductive responses to males, even in the absence of previous sexual experience, when sexually active bucks are used.     

Parental care behavior in the monogamous,sexually dimorphic Madagascar paradise flycatcher: sex differences and the effect of brood size

I studied the parental care behavior of the Madagascar paradise flycatcher Terpsiphone mutata in northwestern Madagascar. I especially focused on feeding, brooding and vigilance behaviors. Feeding rate did not differ between males and females, but females spent more time at the nest than males. Females dedicated their time to brooding, while males perched on the nest and were vigilant. Both parents changed the feeding rate in relation to brood size, so the feeding rate per nestling was not different among nests of different brood size. Duration of brooding by females increased with decreasing brood size, suggesting that the Royama effect, the pattern of lower feeding rate per nestling in larger broods, did not apply in this study. Males spent more time on vigilance than females. Anti-predator vigilance by males should be important for nestling survival given the high predation pressure typical of this population. In conclusion, males provide considerable parental care probably to minimize nestling starvation and to avoid nest predation. My results are not consistent with the general pattern of less parental effort by males in monogamous, sexually dimorphic species.     

The significance of biparental care in the brown bullhead,Ictalurus nebulosus

Synopsis Parental care in the brown bullhead is characterized by variation in the participation of each sex. Most broods are attended by both sexes, but some are attended by a male alone, or rarely, a female alone. Two care-givers were more successful than one alone in fostering offspring survival. However, there was no significant difference between two care-givers and one alone in the proportion of time that broods were unattended. Potential brood predators were chased less frequently by one adult alone than by adults aided by their mate. This difference may be unimportant since two adults simultaneously attended their brood only 19% of the time. Males alone attended their broods a significantly greater proportion of time than did either males or females aided by their mates. This difference suggests that males alone sustain a greater cost of care-giving (starvation and therefore reduced future reproduction) than do males aided by their mates. Thus, males alone may more often leave broods (and not return) than males that are aided in care-giving. The differential success observed may be due to a difference in the likelihood that the male (the principal care-giver) leaves the brood permanently, rather than differences in the quality of care one or two adults provide. I suggest that two care-givers are more successful than one because the net benefits of care-giving exceed the net benefits of leaving for males when aided by their mates.     

Why spacing behavior does not stabilize density in cyclic populations of microtine rodents

Summary There are several published hypotheses that consider spacing behavior to be a significant factor causing the multiannual density fluctuations characteristic of some microtine rodent populations. Recent modeling efforts have concluded, however, that spacing behavior should have a stabilizing rather than a destabilizing effect on population dynamics. Why doesn't spacing behavior stabilize these cyclic populations? We argue that while spacing behavior does have a stabilizing influence on population dynamics by limiting the number of breeding individuals, reproduction continues and population size is not limited in an asymptotic manner. Rather, microtine social organization produces demographic changes within a population that allow density cycles to occur under certain conditions. Using a simulation model, we demonstrate that in a strongly seasonal environment populations with low density dependence in reproduction will cycle whereas populations with high density dependence in reproduction will have relatively stable densities. Given such complicating factors as the annual species nature of microtine rodents, occasionally intense predation, and the tendency for territoriality to break down during the non-breeding season, individuals with low density dependence in reproduction will always be able to invade and eventually dominate populations with high density dependence in reproduction, regardless of the resulting destabilization of population dynamics.     

Pollen competition among Populus deltoides Marsh., P. nigra L. and P. maximowiczii Henry in fertilizing P. deltoides ovules and siring its seed crop

Summary Interspecific pollen competition among Populus deltoides, P. nigra and P. maximowiczii in fertilizing P. deltoides ovules was studied by using a pollen mixture technique, allozymes and leaf morphology. The frequencies of F₁ seedlings of different paternities in pollen-mix crosses showed highly significant (P<0.01) departures from the 11 ratio expected if pollen selection was random. P. deltoides pollen was the most competitive. The mean percentages of F₁ seedlings of P. deltoides paternity in crosses with pollen mixes P. deltodes + P. nigra, P. deltoides + P. maximowiczii, and P. deltoides + P. nigra + P. maximowiczii were 95.0, 92.5, and 84.8, respectively. P. maximowiczii pollen was more competitive than P. nigra pollen, which was at a selective disadvantage. An average of 83.6% of F₁ progenies of the eight crosses with P. nigra + P. maximowiczii pollen showed P. maximowiczii paternity. Also, in four crosses with P. deltoides + P. nigra + P. maximowiczii pollen, the relative proportion of P. deltoides × P. maximowiczii seedlings (13.4%) was higher than that of P. deltoides × P. nigra seedlings (1.8%). Pollen proportions in the pollen mixes and pollen size did not significantly affect the competitive ability of the pollen. The relative pollen competitive ability indicated reproductive affinities among the species.     

Bimorphism in Male Verreaux’s Sifaka in the Kirindy Forest of Madagascar

Male primates in species with pronounced secondary sexual adornments can exhibit reversible or irreversible bimorphism, i.e., striking variation in the degree to which males express the adornments. Verreaux’s sifaka (Propithecus verreauxi verreauxi) use scent marking as a form of communication and exhibit sex differences in scent glands. Some males exhibit a pronounced brown staining around their sternal gland, whereas others do not. We studied morphological and behavioral characteristics of males in 6 social groups in Kirindy Forest, Madagascar, from November 2000 to March 2002 to evaluate the hypotheses that the bimorphism in male sifaka chest status represents alternative mating tactics and is a badge of status. Males are clearly divided into 2 categories: clean and stained chests, with rare, but informative, intermediate males. The chest staining probably results from the males scent marking with their sternal glands, because stained-chested males scent marked significantly more often than clean-chested males. Though sample sizes are small, chest status did not appear to depend on body size. Chest status is reversible and related to dominance rank. In each group, only 1 male, the dominant, was stained-chested, whereas all other (subordinate) males were clean-chested. These findings suggest that stained chests are visual and olfactory signals of dominance rank and that clean chests signal lack of competitive intent. Thus, this bimorphism may reflect alternative mating tactics used by males to maximize their reproductive success based upon their social environment.     

Female-to-male shift of mouthbrooding in a cichlid fish,Tanganicodus irsacae,with notes on breeding habits of two related species in Lake Tanganyika

Synopsis Parental-care patterns and mating systems of three goby-like cichlids in Lake Tanganyika were investigated. In Tanganicodus irsacae females mouthbrooded eggs and small young for about two weeks and then males took over the role for about one week. Field observations of tagged fish suggest that this species is monogamous: a male's home range largely overlapped with that of its mate, while their home ranges were segregated from those of similar-sized consexual adults. Eretmodus cyanostictus also performed female-to-male shift of mouthbrooding and appeared to be monogamous. The third species, Spathodus marlieri, however, exhibited exclusively maternal mouthbrooding. The differences in parental care and mating system among the three species are discussed in relation to their feeding habits, and the pattern of monogamy in the goby-like cichlids is compared with those of other fishes.     

Parental care of kestrels living in stable and varying environmental conditions

We studied parental behaviour of two populations of Eurasian kestrels (Falco tinnunculus) living in environments with different reproductive constraints: a northern Italy population, resident, with relatively long reproductive season and almost stable prey availability, and a western Finland population, long-distance migrant, with short reproductive season and fluctuating availability of main prey (voles). The Finnish population was studied during a low vole availability year. We aimed to test the supposition that in variable environments the few pairs capable of breeding during low prey availability years show high-quality parental care. To estimate the quality of parental care in both Finnish and Italian pairs we measured the difference in the number of prey delivered by the male and by the female within each pair. The parental care, measured by that parameter, was greater among Italian pairs during both the incubation and early nestling periods. However, this was not the case in the late nestling period, when food requirements of offspring are greater. Moreover, the Finnish and Italian pairs produced about the same number of fledglings. Therefore, our data supported the initial supposition and we conclude that the few Finnish pairs capable of breeding in a low vole availability year show parental care similar to Italian pairs.     

Risk of sperm competition does not influence copulatory behavior in the promiscuous meadow vole (<Emphasis Type="Italic">Microtus pennsylvanicus</Emphasis>)

It is not clear whether males in all mammalian species adjust their copulatory behavior when faced with risk of sperm competition (RSC). Previous work on meadow voles, Microtus pennsylvanicus, indicated that males increase their sperm expenditure but not the number of ejaculations in the presence of odors of a conspecific male. The present study follows up on this work and asks whether male meadow voles modify any aspect of their copulatory behavior when they face a RSC. We examined 46 variables of copulatory behavior and found that the presence of odors from a conspecific male did not affect any of these variables. Thus, male meadow voles, unlike some other species of mammals, do not adjust their copulatory behavior when exposed to cues associated with an elevated RSC.     

Copulatory Plug Displacement Evidences Sperm Competition in Lemur catta

Male displacement of copulatory (sperm) plugs from female vaginas provides further evidence for sperm competition in ring-tailed lemurs (Lemur catta), a gregarious prosimian species with a multimale, multifemale mating system. During two mating seasons, I studied two groups of free-ranging ring-tailed lemurs on St. Catherines Island, GA, USA. I observed 22 mating pairs in which males achieved penile intromission. Copulatory plug displacement by males occurred in 9 cases. Plugs were displaced during copulation by male penes upon withdrawl following deep vaginal thrusting. In every case of copulatory plug displacement, the male displacing a plug mated to ejaculation with the estrous female. In a mating system in which females typically mate with more than one male during estrous, often in succession, copulatory plug displacement may function to disrupt or preclude other males' successful insemination of estrous females. The effects of sperm plug displacement on paternity in Lemur catta are unknown, as no study had heretofore documented copulatory plug displacement in this species. The first-male mating advantage suggested for Lemur catta should be re-evaluated where mating order is known, and copulatory plug displacement during mating, or lack thereof, is identified. Because there is a tendency for first-mating males to mate-guard for longer periods of time in Lemur catta, the latency period between the first mate's ejaculation and that of subsequent mates may be an important determinant of male fertilization success.     

Reproductive tactics of the Asian cichlids of the genusEtroplus in Sri Lanka

Synopsis The Asian cichlids,Etroplus maculatus (the orange chromide) and E. suratensis (the green chromide) in Sri Lanka reproduce twice during the year when water conditions are favorable for nest construction and maintaining visual contact with offspring. These are the drier premonsoonal and monsoonal seasons when water turbidity decreased and salinity increased. When breeding in isolation orange chromide pairs selected dense vegetation where nests were camouflaged. During the peak breeding cycle (July) orange chromide pairs selected sparse vegetation for nesting as a compromise between survival of young and availability of adult food. These areas were also occupied by foraging non-breeding conspecifics which increased the threat of cannibalism of offspring. Under these pressures most orange chromides nested in colonies which helped decrease both actual and attempted cannibalism. The sympatric green chromide does not forage during nesting and nest site selection was determined mainly by factors favoring offspring survival. Biparental care is exhibited by both species. One member of an orange chromide pair stands guard over offspring while the other leaves the territory to forage — their roles are reversed every few minutes. The total parental investment is equivalent to the full investment of a single parent. Monogamy appears to be maintained by their metabolic constraints. In the green chromide both parents are vigilant over offspring and neither forages thus spending twice as much time in parental investment.     

Ultrastructural examination of the insemination reaction in Drosophila

The insemination reaction is a swelling of the female vagina caused by the male ejaculate. This postmating phenomenon is common among species in the genus Drosophila. It could act as a plug securing male paternity. It is not clear, however, what benefits it provides to the female. The structure formed in the female vagina is expelled in some species and disappears gradually in others suggesting different phenomena. Based on ultrastructural examination of the vaginal contents of five Drosophila species (D. mettleri, D. nigrospiracula, D. melanogaster, D. mojavensis, and D. hexastigma), we propose three terms to describe these vaginal structures: the sperm sac, the mating plug, and the true insemination reaction. Each term describes a distinct structure associated with a specific female postmating behavior. This study questions the concept of the insemination reaction as a single phenomenon and discusses its possible functions from an evolutionary perspective.