Paternity in the Australian brush-turkey, Alectura lathami, a megapode bird with uniparental male care

Male Australian brush-turkeys, Alectura lathami, provide allparental care by building and tending large incubation mounds.Females visit and lay eggs in the mounds of several males sequentially,but they provide no parental care after laying. Because malesand females meet only briefly at mounds to copulate and lay,males have no obvious means of ensuring paternity. I used DNAfingerprinting techniques to determine paternity for 65 brush-turkeychicks. Eighteen chicks (27.7%) did not match the mound-tendingmale. Some of these paternity exclusions were evidently causedby females switching rapidly from one mound to another, butthe majority (23.1% of eggs) appeared to result from femalescopulating with males other than the one in whose mound theywere currently laying. However, the frequency of these copulations(43%) was much lower than the estimated frequency with whichthey fertilized eggs, perhaps because their timing during theovulatory cycle differed relative to most other copulations.The percentage of eggs excluded in paternity analyses rangedfrom 20.0% to 43.8% for individual males but did not appearto affect male parental care. Several factors favor male parentalcare regardless of paternity. Males can accommodate eggs fromseveral females in one mound, which increases the opportunitiesfor additional matings without increasing the cost of parentalcare. In addition, paternity appears to be unpredictable andhard to assess, and a facultative reduction in care would bedifficult without abandoning a mound entirely.     

Selection of incubation mound sites by the Australian Brush-turkey Alectura lathami

Environmental characteristics associated with siting of the incubation mounds of the megapode, the Australian Brush-turkey Alectura lathami were investigated in three rainforest locations in southeast Queensland. Seventeen environmental variables were measured for 52 mounds and 137 non-mound sites. Most mounds were constructed on top of old mounds which had not been used during the previous year. A Discriminant Function Analysis obtained a clear separation between mound and non-mound sites, based primarily on three variables: (1) the number of saplings growing within the mound or centre of the site: this was considered to be a result of the germination of seeds raked in earlier years; (2) the proportion of Lantana camara thicket occupying the site. Approximately half of the mounds constructed during a season were abandoned by their constructor due to expulsion by a neighbouring male. Brush-turkeys could more easily establish mounds next to thickets since the efficiency of defence by other males was reduced in these areas; and (3) the openness of the canopy. Areas with greater canopy cover provided two important mound resources: a good supply of leaf litter, and protection of the mound from dessication.     

Burrow architecture and digging activity in the Cape dune mole rat

While females are traditionally thought to invest more time and energy into parental care than males, males often invest more resources into searching and displaying for mates, obtaining mates and in male–male conflict. Solitary subterranean mammals perform these activities in a particularly challenging niche, necessitating energetically expensive burrowing to both search for mates and forage for food. This restriction presumably affects males more than females as the former are thought to dig longer tunnels that cover greater distances to search for females. We excavated burrow systems of male and female Cape dune mole rats Bathyergus suillus the, largest truly subterranean mammal, to investigate whether male burrows differ from those of females in ways that reflect mate searching by males. We consider burrow architecture (length, internal dimensions, fractal dimension of tunnel systems, number of nesting chambers and mole mounds on the surface) in relation to mating strategy. Males excavated significantly longer burrow systems with higher fractal dimensions and larger burrow areas than females. Male burrow systems were also significantly farther from one another than females were from other females' burrow systems. However, no sex differences were evident in tunnel cross-sectional area, mass of soil excavated per mound, number of mounds produced per unit burrow length or mass of soil excavated per burrow system. Hence, while males may use their habitat differently from females, they do not appear to differ in the dimensions of the tunnels they create. Thus, exploration and use of the habitat differs between the sexes, which may be a consequence of sex differences in mating behaviour and greater demands for food.     

Components of care: are they honest signals of parental care quality?

It has been argued that male parental care provides direct benefits to females and therefore should be under sexual selection. Given this, we expect signals that honestly indicate the quality of care to be favoured by selection. One such potential signal is care itself. Fish have several features that make them excellent model systems for studying the evolution and dynamics of parental care. We use the flagfish, Jordanella floridae , as a model to evaluate these general ideas. Males of this species guard, clean and fan empty nests and then eggs. Females prefer males that fan more (1) before spawning and (2) when eggs are newly received. When single males and females were paired, males that fanned and visited their nests more prior to spawning were more likely to be mated. Furthermore, among successful males, rates of fanning in the first day after spawning were correlated with the number of eggs received in the future (but not current egg numbers). We then considered whether these two putative signals were correlated and whether males that fan more in these contexts actually have higher egg survivorship. We found no correlation between nest fanning rates before and after spawning and neither 'signal' was predictive of variation in egg survivorship among mated males. We further considered whether pre‐spawning fanning rates were predictive of hatching success in an experiment in which single males were allowed to establish nests and provided eggs. We found little evidence that fanning is an honest signal of care quality and discuss alternative explanations. In particular, we discuss patterns of care elaboration in light of our results.     

Effects of northern pike on patterns of nest use and resproductive behavior of male fathead minnows in a boreal lake

We conducted a two-part study to assets predator avoidance byreproductive male fathead minnows (Pimephales promelos) subjectedto predation threat from northern pike (Esox lucius). First,we determined if patterns of nest use by egg-guarding male minnowsin a boreal lake were related to pike densities. We samplednorthern pike and identified four areas of "high pike-density"and three areas of "low pike-density." We censused natural nestsand placed nest boards in these areas. We found eggs on naturalnests more frequently in areas with low densities of pike thanin areas with high densities of pike. However, we could notfully explain the distribution of nests by predation risk. Second,we evaluated the behavioral response of egg-guarding males toa control stimulus (a piece of wood) or a live pike in a wirecage. We used time to return to the nest after a stimulus asa measure of risk taking. Males took different amounts of riskbased on predation threat; males in the predator treatment tooklonger to return to their nests than control males. Risk takingwas not related to the number or age of the eggs but to distanceto nearest egg-guarding neighbor; males with close neighborsreturned sooner than more isolated males. Males in the predatortreatment had lower total activity and egg rubbing than controlmales after they returned to their nests. We conclude that malefathead minnows altered their reproductive behavior in waysthat reduced predation risk, but the cost of predator avoidancemight include egg predation, lost mating opportunities, or usurpationof nests     

Are males with more attractive resources more selective in their mate preferences? A test in a polygynous species

In many polygynous species, males typically direct more intensecourtship toward more fecund females. Here we examined thisbehavior in relation to the attractiveness of a male's resource.We used the territorial polygynous beaugregory damselfish (Stegastesleucostictus) and manipulated the quality of male breeding territorieswith two types of artificial sites. We also investigated variablenatural breeding territories. Previous studies have shown thatthese different breeding sites were of different qualities,as judged by the number of eggs accrued by the defending male.Males on all three types of breeding sites did court females,and males using the highest quality sites exhibited significantlyhigher courtship intensity. However, only the group of maleson the highest quality site-type modulated their courtship intensityto female quality (i.e., female size). This indicates that males requiredsome minimal level of resource attractiveness (i.e., a threshold) beforethey exhibited mate preferences based on female quality. Further differencesin the resource attractiveness for males defending the high-qualityartificial sites were not related to differences in courtship behavior.     

Ventilation of termite mounds: new results require a new model

In 1955, Lüscher proposed a ventilation mechanism forcathedral-shaped termite mounds to exchange respiratory gases. This mechanism was generally accepted, although it had neverbeen tested critically. We tested this mechanism by investigatingtemperatures, CO₂ concentrations, and air currents in and aroundtwo types of Macrotermes bellicosus mounds: cathedral-shapedmounds with many ridges and thin walls located in the savannaand dome-shaped mounds without ridges and with thick wallsin the forest. These two mound shapes have two different mechanismsof ventilation, depending on the environmental temperature.In the savanna during the day, sun heats the air in the peripheralair channels inside the ridges of the mound above the centralnest temperatures and produces a temperature gradient in theperipheral air channels with decreased temperatures at thetop of the mound. This temperature gradient leads to convectioncurrents with air rising inside the air channels of the ridgesto the top of the mound, meanwhile exchanging CO₂. In contrast,in the savanna during the night and generally in the forest,the temperatures inside the air channels are lower than thoseof the central nest, and no air currents rising upward insidethe air channels were detected. The CO₂ concentrations in theair channels of savanna mounds at night and forest mounds ingeneral were higher than during the day in the savanna. Therefore,our data do not support Lüscher's proposed mechanism.     

Predation of incubator birds (Megapodius freycinet) by Komodo dragons (Varanus komodoensis)

A short study was made of the nesting habits of a species of megapode, Megapodius freycinet on the island of Komodo. The birds were found to build giant nesting mounds (0.95 × 7.15 m) in scrub and woodland close to the seashore or alongside river beds; 23 active and 19 abandoned mounds were located in an area of 250 ha.
The mounds were apparently added to from year to year, and consisted of dry sand and soil, plus leaves and sticks in most cases. The upper parts of the mounds were invariably exposed to the direct rays of the sun; the temperature within the active mounds was 33–37° C and slightly less in mounds not in use by the birds. According to the local people the nesting season of the megapodes extended from late August to November, and the chicks were hatched before the arrival of the rains in December. Up to four eggs had been recovered from a single mound.
The active mounds were all spaced apart by at least a 100 m, and the closest ones were usually separated by a river bed. This pattern suggested the species was territorial. One or two birds were often seen scraping at a mound or feeding close by, and their behaviour was recorded.
The Komodo dragon, Varanus komodoensis , was a frequent visitor to the nesting mounds of the megapodes, and was able to burrow into them and steal the eggs.     

Why do the mounds of Mus spicilegus vary so much in size and composition?

Mound-building mice (Mus spicilegus) construct large and often enormous mounds, which may have several functions, including food storage, thermoregulation, and protection from predators. We examined variation in mound size and composition to help determine their adaptive significance. Like previous studies, we found much variation in mound size within and between geographic populations, and the main differences were between habitats: we found up to a 700-fold variation in mound size between corn fields and harvested grain fields. Mice mainly utilized three plants (Amaranthus spp., Chenopdium spp. and Setaria spp.) as building materials, and the availability of these may affect mound size. We found a positive correlation between the availability of plant material in the surrounding of the mound and mound size. Surprisingly, mound size was independent of the number of inhabitants within the mounds, which indicates that larger mounds are not constructed to shelter more individuals. Future studies need to examine how variation in mound size and composition affects overwinter survival, and also which individuals contribute to mound construction.     

Community response to experimental soil disturbance in a midsuccessional,abandoned pasture

To determine whether soil disturbance by digging and burrowing mammals altered community structure and the rate of succession in a midsuccessional abandoned pasture, species richness, composition and relative abundance were monitored over a two year period both on and off artificially created earth mounds (100, 900, 8100 cm²). Mean species richness increased by up to two species per small mound (100 cm²) and by up to four species per large mound (8100 cm²). However, increased species richness was evident for less than two years. Initially, up to sixteen of the twenty species present occurred more often on earth mounds than off mounds, with two of these species found only on large mounds (8100 cm²). After two years, there was little or no significant difference in species composition and relative frequency on and off earth mounds. Experimental soil disturbance temporarily altered community structure simply by increasing space available for colonization since light, nutrient and water supply did not increase significantly on mounds. Soil disturbance can increase species richness and change species' relative frequency in disturbances as small as 100 cm² but such changes were likely too small and short lived to alter permanently the structure and rate of succession in the abandoned pasture studied here.     

Spatial variability and abiotic determinants of termite mounds throughout a savanna catchment

Termite mounds contribute to the spatial heterogeneity of ecological processes in many savannas, but the underlying patterns and determinants of mound distributions remain poorly understood. Using the Carnegie Airborne Observatory (CAO), we mapped the distribution of termite mounds across a rainfall gradient within a river catchment (～ 27 000 ha) of the Kruger National Park, South Africa. We assessed how different factors were associated with the distribution and height of termite mounds at three spatial scales: the entire catchment, among three broad vegetation types, and on individual hillslope crests. Abiotic factors such as the underlying geology and mean annual precipitation shaped mound densities at broad scales, while local hillslope morphology strongly influenced mound distribution at finer scales, emphasising the importance of spatial scale when assessing mound densities. Fire return period had no apparent association with mound densities or height. Mound density averaged 0.46 mounds ha^?1, and exhibited a clustered pattern throughout the landscape, occurring at relatively high densities (up to 2 mounds ha^?1) on crests, which are nutrient‐poor elements of the landscape. Mounds exhibited significant over‐dispersion (even spacing) at scales below 60 m so that evenly spaced aggregations of termite mounds are embedded within a landscape of varying mound densities. The tallest mounds were found in dry savanna (500 mm yr^?1) and were positively correlated with mound density, suggesting that dry granitic savannas are ideal habitat for mound‐building termites. Mound activity status also varied significantly across the rainfall gradient, with a higher proportion of active (live) mounds in the drier sites. The differential spacing of mounds across landscapes provides essential nutrient hotspots in crest locations, potentially sustaining species that would otherwise not persist. The contribution to biodiversity and ecosystem functioning that mounds provide is not uniform throughout landscapes, but varies considerably with spatial scale and context.     

Effect of some biological factors on soil variability in the tropics

Summary Observations of a selected area in the forest zone of south western Nigeria indicate that the distribution ofMacrotermes bellicosus (Smeathman) andMacrotermes subhyalinus (Rambur) mounds are mainly affected by the drainage conditions of the soils along a toposequence. Higher number of mounds are observed on the better drained soils, ranging from 35.4 to 17.2 mounds/ha covering an area of respectively 0.84 and 0.10 per cent. The shallow and poorly drained soils show lower mound density (<1 mound/ha). The mound soil bears close resemblance to the subsoil, has higher clay content, lower pH, organic C, CEC and extractable P, but higher P retentio than the adjacent surface soil. Growth of maize (Zea mays L.) and soybean (Glycine max. (L.) Merrill) was poorer in the mound soil than in the surface soil. The problem of increased soil variability when large number of mounds are encountered in the field is discussed.     

The role of mounds in promoting water-exchange in the egg-tending burrows of monogamous goby, Valenciennea longipinnis (Lay et Bennett)

Valenciennea longipinnis spawns monogamously in a burrow. After spawning, the paired female constructs a conspicuous mound on the burrow by carrying and piling up substratum-derived materials while the male tends eggs in the burrow until hatching occurs. In this study, the mounds of V. longipinnis were tested in the field to confirm their function of promoting water-exchange in the burrow, and their ecological role was examined in relation to egg care by the male. The mound of V.longipinnis promoted water-exchange in the burrow, contributing to the provision of external oxygenated sea water into the burrow. Therefore, dissolved oxygen (DO) concentrations in the burrow with a mound were significantly higher than those without a mound. Although male egg-tending behaviour (e.g., fanning) may also promote water-exchange in the burrow, the water-exchange appeared to depend mainly on the hydrodynamic effect. Removals of the mound and paired female on the day of spawning led to high rates of egg-desertion by males. Since the frequency and time of fanning increase with a decrease of DO concentration in the burrow, the egg-desertion may result from an increased parental cost to males due to the decrease of water-exchange without a mound. This was supported by the fact that the DO concentrations on the day after mound removal were significantly lower in the egg-deserted burrows (measured before desertions) than in burrows not deserted by the male. Moreover, removals of paired females only also led to higher desertion rates. After removal of the female, the mound gradually collapsed by wave action and other factors, and the surface of the mound was covered with planktonic materials. Such a mound of poor quality may provide little water-exchange, which may lead to the egg-desertion by males. These results indicated that mound maintenance by females during the egg-tending period has an important role in the success of parental care by males.     

Hydrophobins Sc3 and Sc4 gene expression in mounds, fruiting bodies and vegetative hyphae of Schizophyllum commune.

An abnormal growth form called mound has been hypothesized to be a neoplasm in the filamentous fungus Schizophyllum commune. An alternative hypothesis is that mounds represent some unusual developmental form in the fruiting body morphogenetic pathway. Hydrophobin proteins have been found in fruiting bodies where they line the surface of gas exchange pores and function to keep the pores hydrophobic. To further determine possible relationships between mounds and fruiting bodies, mound tissue was examined for gas exchange pores and the presence of hydrophobins. Cryoscanning electron microscopic images revealed the presence of channels in mound tissue and presumptive hydrophobin rodlets similar to the air channels in fruiting bodies. Hydrophobin gene expression was also measured in mound tissue using quantitative real-time PCR and showed both monokaryotic and dikaryotic mound tissue exhibited high expression of the dikaryotic specific Sc4 hydrophobin gene. In contrast, Sc4 hydrophobin expression was barely detectable in monokaryotic fruiting bodies. The expression of Sc4 hydrophobin genes in mounds suggests mound development uses this aspect of the dikaryotic fruiting developmental pathway.     

Hydrophobins Sc3 and Sc4 gene expression in mounds, fruiting bodies and vegetative hyphae of Schizophyllum commune

An abnormal growth form called mound has been hypothesized to be a neoplasm in the filamentous fungus Schizophyllum commune. An alternative hypothesis is that mounds represent some unusual developmental form in the fruiting body morphogenetic pathway. Hydrophobin proteins have been found in fruiting bodies where they line the surface of gas exchange pores and function to keep the pores hydrophobic. To further determine possible relationships between mounds and fruiting bodies, mound tissue was examined for gas exchange pores and the presence of hydrophobins. Cryoscanning electron microscopic images revealed the presence of channels in mound tissue and presumptive hydrophobin rodlets similar to the air channels in fruiting bodies. Hydrophobin gene expression was also measured in mound tissue using quantitative real-time PCR and showed both monokaryotic and dikaryotic mound tissue exhibited high expression of the dikaryotic specific Sc4 hydrophobin gene. In contrast, Sc4 hydrophobin expression was barely detectable in monokaryotic fruiting bodies. The expression of Sc4 hydrophobin genes in mounds suggests mound development uses this aspect of the dikaryotic fruiting developmental pathway.     

Social structure of the mound-building mouse Mus spicilegus revealed by genetic analysis with microsatellites

The Mound-building mouse Mus spicilegus possesses a unique behaviour amongst mice. It constructs large earthen mounds and associated nesting chambers which serve to store food for immature individuals during the winter nesting period. We have used genetic analysis of four autosomal and four X-linked microsatellite loci to determine relationships between individuals inhabiting 40 mounds in Bulgaria. We show that, in almost all cases, individuals in a mound are the product of multiple parentage. We estimate the minimum number of males and female parents contributing offspring to each mound and demonstrate that at least two male and two female parents contribute offspring to a minimum of seven mounds. Analyses of relatedness coefficients and allele sharing values demonstrate that parents of different sibships within mounds are more related than if they had been chosen at random from the population and suggest that it is the female parents that contribute this excess relatedness. These results suggest that the mechanism by which individuals congregate to build mounds is kin-based and that the evolution of mound building and communal nesting in M. spicilegus is due in part to kin selection. This study represents a novel approach to the study of mammalian behavioural ecology. We have used a genetic dataset to construct an outline of social structure in the absence of behavioural data. These inferences can now be used to direct further work on this species.     

Termites (Isoptera) as secondary occupants in mounds of Macrotermes michaelseni (Sj?stedt) in Kenya

Termites were found as secondary occupants in mounds of Macrotermes michaelseni (Sj?stedt) in Kajiado District, in the Rift Valley Province of Kenya. Data are presented from 34 mounds that were completely demolished and extracted to analyse the population of the primary nest. Secondary occupants were found in 29 of the 34 mounds, averaging 2.4 species per occupied mound (or 2.1 species per mound overall). The termites belonged to nine species, three of which were not found anywhere else in the area of study. The frequency of occurrence of the different species and their degree of association with the host mounds are examined and discussed.     

Filial cannibalism in an assassin bug

Filial cannibalism has been described in many fish species with male parental care, and has typically been explained as a response to high energetic costs of brood defence and decreased feeding opportunities during the period of care. We investigated filial cannibalism in an insect, the assassin bug Rhinocoris tristis. In this species, males guard eggs of a number of females, cannibalizing some of their offspring within the brood. We monitored guarding males in both the field and the laboratory. Males typically ate eggs around the periphery of the brood, which were those most likely to have been parasitized by wasps. However, cannibalism persisted in the laboratory in the absence of parasites, and the number of cannibalized eggs was related to the length of care and overall brood size, suggesting that males use eggs as an alternative source of food. This conclusion was further supported by the fact that males in the field did not lose weight while guarding, despite being unable to forage efficiently while caring. Males were also observed to adopt broods, but in a laboratory experiment did not eat more eggs from adopted than from their own broods.     

Behavioural and ecological aspects of nuptial flights of the termitophilous phorids Termitophilomyia zimbraunsi and Mesopathusa modesta (Diptera: Phoridae) in Zimbabwe

Nuptial flights of Termitophilomyia zimbraunsi were observed over three years (1990-1993) near the open chimney of an Odontotermes transvaalensis mound in Harare, Zimbabwe. Copulating phoretic pairs landed on the chimney; males quickly disengaged and flew away whilst females ran inside the vent and tore off their wing membranes. Agitated termite workers and soldiers swarmed the rim in a partially successful physical attempt to expel the phorids; the termites appeared to anticipate the threat of invasion. After acquiring the scent of the nest, female phorids ran underground to lay their large single eggs, probably in the hosts'fungus gardens. Nuptial flights occurred throughout the year but only in the early mornings, peaking in the hot dry season and being least in the hot wet season. The flights were highly temperature dependent (15.0-17.9 °), usually occurring at relatively low light intensities but across a wide range of humidities. Phoresy and sexual dimorphism in wings, mouthparts, food and division of labour in T. zimbraunsi are discussed, as is its termitophilous habit and the responses of termites to it.
Although the females of Mesopathusa modesta are apterous, nuptial flight behaviour is very similar to that of T. zimbraunsi. Neither phorid is host-specific, both species having been found on both Odontotermes latericius and O. transvaalensis mounds, although M. modesta is more common on O. latericius mounds.     

Alternative mate-finding tactics in a non-territorial damselfly (Odonata: Coenagrionidae)

Males of the non-territorial damselfly Enallagma hageni have two alternative tactics for finding mates: (1) they search the banks of the pond for unmated females (searching tactic), or (2) wait at oviposition sites for females that resurface prematurely from underwater oviposition (waiting tactic). Although the searching tactic yielded more fertilizations than the waiting tactic, for time invested, the waiting tactic became increasingly successful later in the reproductive season due to changes in female oviposition behaviour. The two tactics can be maintained in the population because males can mate by the waiting tactic during the afternoon when few females are available to searchers. Among males visiting the breeding site an equal number of times, males mating by a mixture of tactics were as successful as males mating only by the main tactic. Because marked males were found to use both tactics, these behaviours are interpreted as evidence of behavioural plasticity within individuals, representing one conditional evolutionary strategy.