Lifetime reproductive success in the spotted sandpiper (Actitis macularia) : sex differences and variance components

Summary Lifetime reproductive success (LRS) of male (N=103) and female (N= 66) spotted sandpipers (Actitis macularia) was studied for 13 years of a 17-year study at Little Pelican Island, Leech Lake, Minnesota. There was no sex difference in longevity, but females had significantly more mates, eggs, chicks, fledged young, and young returning in subsequent years than did males. Variance in LRS was partitioned into five life-history components: longevity (L), mates per year (M), eggs per mate (E), proportion eggs hatched (H), and proportion of chicks fledged (F). For both sexes, F accounted for the greatest proportion of variance in LRS (males, 43%; females, 47%), followed by L (males, 26%; females, 43%) and H (males, 21%; females, 28%). Positive covariance between H and F was consistent with predator-caused clutch and brood loss. Contrary to our expectations, males had a higher coefficient of variation in reproductive success than did females. This was because males were relatively more likely than females to produce no young.Offprint requests to: L.W. Oring at the current address     

Effects of changes in water salinity upon exercise and cardiac performance in the European seabass (<Emphasis Type="Italic">Dicentrarchus labrax</Emphasis>)

The European seabass is an active euryhaline teleost that migrates and forages in waters of widely differing salinities. Oxygen uptake (M_O2) was measured in seabass (average mass and forklength 510 g and 34 cm, respectively) during exercise at incremental swimming speeds in a tunnel respirometer in seawater (SW) at a salinity of 30 and temperature of 14°C, and their maximal sustainable (critical) swimming speed (U_crit) determined. Cardiac output (Q) was measured via an ultrasound flow probe on their ventral aorta. The fish were then exposed to acute reductions in water salinity, to either SW (control), 10, 5, or freshwater (FW, 0), and their exercise and cardiac performance measured again, 18 h later. Seabass were also acclimated to FW for 3 weeks, and then their exercise performance measured before and at 18 h after acute exposure to SW at 30. In SW, seabass exhibited an exponential increase in M_O2 and Q with increasing swimming speed, to a maximum M_O2 of 339±17 mg kg^–1 h^–1 and maximum Q of 52.0±1.9 ml min^–1 kg^–1 (mean±1 SEM; n=19). Both M_O2 and Q exhibited signs of a plateau as the fish approached a U_crit of 2.25±0.08 bodylengths s^–1. Increases in Q during exercise were almost exclusively due to increased heart rate rather than ventricular stroke volume. There were no significant effects of the changes in salinity upon M_O2 during exercise, U_crit or cardiac performance. This was linked to an exceptional capacity to maintain plasma osmolality and tissue water content unchanged following all salinity challenges. This extraordinary adaptation would allow the seabass to maintain skeletal and cardiac muscle function while migrating through waters of widely differing salinities.Communicated by S.A. Poulet, Roscoff     

Allozyme and colour differences between two sibling species of the heteronemertean Lineus torquatus from the Sea of Japan

Brown and cherry colour morphs of the heteronemertean Lineus torquatus Coe, 1901 from the Vostok Bay of the Sea of Japan were compared using allele frequencies at 18 isozyme loci. These morphs shared no common alleles at 9 of the 18 loci. The observed genetic differences are strong evidence that the brown and cherry colour morphs, which occur sympatrically, are different species. Nei's genetic distance between these species (D=1.135) is considerably higher than typical D values for congeneric animal species. As with much previous work on nemerteans, intraspecific allozymic variability in both species was high, with mean expected heterozygosity values 0.286±0.049 and 0.159±0.051 for L. torquatus (brown) and L. torquatus (cherry) species, respectively.     

On mathematical simulation of a pelagic ecosystem in tropical waters of the ocean

Consideration is given to a finite-difference model of functioning of a pelagic community inhabiting the upper 0 to 200 m layer of the tropical waters of the ocean. The model was developed on the basis of original data, obtained mainly on the 44th cruise of the R.V. Vityaz in the western Equatorial Pacific Ocean. Changes in the system are studied in time (up to 100 days, with 1 day intervals) and in depth (from surface to 200 m depth, with 10 m intervals). The state of the system in the upwelling zone, assumed to be characterized by a homogeneous vertical distribution of all its elements, is regarded as initial. The system then develops with time while it moves with the water flow. The model was realized on a BESM-3M computor. The computations yielded a pattern of changes in time of the biomass of the elements (expressed in calories), and a pattern of their vertical distribution at different instants. The vertical distribution pattern of the major elements of the model shows a rather close correlation with their observed distribution in the ocean at the corresponding instant.     

A role forAmanita muscaria L. in the circulation of cadmium and vanadium in a non-polluted woodland

The sporophore of the fungusAmanita muscaria L. contains greatiy elevated levels of cadmium (29.9 g g^–1 dwt) and vanadium (344.9 g g^–1 dwt) in comparison with the soil in a birch woodland (total (HNO₃-extractabie Cd 0.4 g g^–1 dwt, V 11.7 g g^–1 dwt). The significance of this remarkable concentration of normally rare and dispersed elements in terms of their circulation in the woodland has been investigated. Both elements are released from sporophore tissue in a form which can be taken up by a test plant (lettuce), cultivated in the woodland soil amended with different quantities of sporophore tissue, Cadmium levels in all plant tissues were elevated in comparison to the non-amended controls; only root vanadium levels responded to the amendment of the soil. The results are discussed in terms of their significance for the natural cycling of both elements. It is calculated that an abundant population of sporophores could circulate 1.4% of the total cadmium and 0.65% of the total vanadium pool found in the litter layer and 0–5 cm soil horizon in the sampled woodland over a period of 14 days (mean life span of a sporophore).     

The composition and biosynthesis of lipids in Thysanoessa raschi from the Clyde Estuary,Scotland

The lipid composition and biosynthesising activity of Thysanoessa raschi collected from the Clyde Estuary, Scotland, in May 1981 were examined. Triacylglycerols were the major lipid class present, although 16.7% of the total lipid were wax esters in which phytol was the dominant fatty alcohol. The thoracic contents (hepatopancreas) of the krill were capable of biosynthesising lipids in vitro from various labelled substrates. Radioactivity from [1-¹⁴C] palmitic acid was incorporated into lipids in the order phospholipids>triacylglycerols>wax esters; the bulk of the radioactivity was present in all cases in the fatty acyl moieties of the lipids. [U-¹⁴C] glucose labelled lipids in the order phospholipids>triacylglycerols>free fatty acids> was esters; in the first two lipids the radioactivity was mainly in the glycerol moieties, whereas in was esters it was solely in the fatty acyl moieties. The extent of labelling of these lipids from [U-¹⁴C] alanine was less than that from [U-¹⁴C] glucose, but the pattern of labelling was generally similar. More than 90% of the radioactivity incorporated into total lipid from ³H₂O was present in free fatty acids from which it was calculated that the hepatopancreas of T. raschi can synthesise 2.5 g of fatty acid per hour at 15°C. This value is approximately three times lower than that previously determined for T. inermis from Balsfjorden, northern Norway. The results are discussed in terms of the sources of the dietary lipids of krill and the role of endogenous biosynthesis in contributing to its lipid reserves.     

Correlates of male mating success in two species of Anthidium bees (Hymenoptera : Megachilidae)

Summary Field observations were made on the mating behavior of two congeneric species of solitary bees, Anthidium porterae in an arid grassland and A. palliventre in a coastal sand dune habitat. Males of both species exhibited resource defence polygyny and defended hostplants to gain access to females foraging for nectar and pollen. The mating frequencies of marked and measured resident (territorial) males were monitored during periods of continuous observation, following which measurements of territory size and floral resources were obtained. Mating success of A. palliventre males was strongly influenced by territory characteristics: Males that defended small areas with a few rich hostplant patches mated more often than males that held larger territories containing many hostplant patches of low floral density. Large males generally held high-quality sites and thus had a mating advantage over smaller individuals. In A. porterae, on the other hand, male mating success was unrelated to any measure of territory quality. Copulation frequency and male size were positively correlated, however, apparently due to the increased ability of large males to seize and hold females for mating. The two species also differed in the incidence of non-territorial, sneaky males. While absent in A. palliventre, sneaky males accounted for 12% of all mating observed in A. porterae. Males of A. porterae that displayed sneaky tactics mated, on average, as often as resident males. Offprint requests to: E.M. Villalobos     

Spatial mesoscale patterns of West Pacific picophytoplankton as analyzed by flow cytometry: their contribution to subsurface chlorophyll maxima

Vertical distributions of picophytoplankton (ppp) (<2 m) were studied by ship-board flow cytometry during two cruises in Western Pacific waters to Palau and to Australia in 1990. Weak red-fluorescing small ppp, supposed to be free-living prochlorophytes (Chisholm et al. 1988), were abundant in the area surveyed. These ppp, designated the prochlorophytes, were abundant in the surface waters (>10⁴ cells ml^-1) at the northern region (27°03N; 7°11N) in November, whereas in December at the southern tropical stations (0°23.54S; 9°20.30S; 13°50.6S), they formed subsurface maximum layers (>10⁵ cells ml^-1) on a nitracline at a depth of 3.5 to 5.4% surface irradiation. Their fluorescence intensity increased with depth below 10% surface irradiation. The prochlorophytes at a depth of 1% surface irradiation had ten times higher fluorescence than those at the surface layer. The total fluorescence intensity of the prochlorophytes accounted for 32 to 63% of the sum of the total fluorescence intensity of all fluorescing phytoplankton detected at subsurface chlorophyll maxima in the tropical area. These results suggest that distribution of the prochlorophytes is greatly affected by nitracline and by light intensity and that their chlorophyll is a major contributor to the subsurface chlorophyll maximum in the pelagic West Pacific Ocean.     

Skin papillae as possible electroreceptors in the deep-sea eel Cyema atrum (Cyemidae: Anguilloidei)

Surface structure and internal organization of prominent skin papillae in the mesopelagic eel Cyema atrum have been investigated by scanning and transmission electron microscopy. Each papilla is club-shaped and possesses a pore at its distal end. A central rod, filled with cellular strands and surrounded by a 30 to 40 m wide matrix of clear material, runs from the tip to the base of the papilla. A cluster of cells, located 60 m below the surface of the skin, appears to be connected with the lateral line nerve. The lack of stereo- and kinocilia on the one hand, and the presence of a central rod and a terminal pore on the other, demonstrate that the cutaneous papillae in C. atrum cannot be regarded as typical lateral-line neuromasts. They share several features with electroreceptors, known from other fishes. It is postulated that C. atrum and other deep-sea species with similar skin papillae have been able to partially substitute vision and other senses useful for orientation by electroreception.     

Biology of penaeid prawns in the Suez Canal lakes

A study was made from January 1988 to March 1989 of the penaid prawns in the Great Bitter Lake and Lake Timsah located in the central part of the Suez Canal. Two species of Red Sea origin were investigated,Metapenaeus stebbingi andTrachypenaeus curvirostris; the former is by far the commoner. Both species displayed seasonal breeding over the period April to October, with peaks in the early and late parts of the season.M. stebbingi had two main cohorts each year. A spring-spawned cohort survived 16 mo, and attained modal sizes of 23 mm carapare length (CL) for females and 16 to 17 mm CL for males. The autumn-spawned cohort survived 13 mo, and reached modal sizes of 20 mm and 15 mm CL for males and females, respectively. The reasons for the success ofM. stebbingi in the Canal lakes, in contrast to other Lessepsian penaeids which have flourished in the Mediterranean Sea, have been considered. The main factors are that it is essentially a shallow-water species, with excellent tolerance of salinity variations and adverse environments. However, the Canal lake populations have smaller maximum sizes, and mature earlier, than those in open seas.     

Selective males and ardent females in pipefishes

Summary In the pipefishes Syngnathus typhle and Nerophis ophidion, males have been shown to limit female reproductive rate, and females to compete for access to males. Hence, these species fit the criteria for sex-role reversal. Males brood the eggs and provide the offspring with nutrients, oxygen and an osmoregulated environment. Moreover, in S. typhle both sexes prefer a larger mate when given a choice. Sexual selection theory predicts that males should be more choosy than females, and that was experimentally demonstrated in this study. We predicted that S. typhle males should be less eager to copulate than S. typhle females with an unattractive (i.e. small) mate. We measured eagerness as the time from the start of the experiment until copulation occurred. Males with unattractive partners took significantly longer to copulate than females with unattractive partners. Moreover, females invariably initiated the courtship dance, and resumed it quicker after copulation than did the males, again suggesting reproductive hesitation in males. Neither male nor female size per se was correlated with time until copulation. In N. ophidion, where we have previously shown that males prefer larger to smaller females, we found that females did not select males with regard to size. Our results are consistent both with earlier findings (males limit female reproduction and females compete for males) and with operational sex ratios in nature: in seven annual field samples in June, the numbers of S. typhle females with ripe eggs always significantly exceeded numbers of receptive males. Hence, the potential cost of being choosy in terms of lost matings is much higher in females than in males. In conclusion, S. typhle females were somewhat choosy, but less so than males, whereas N. ophidion females were not choosy at all. Correspondence to: A. Berglund     

Bat serenades—complex courtship songs of the sac-winged bat (<Emphasis Type="Italic">Saccopteryx bilineata</Emphasis>)

Vocalisations of many songbirds, anurans, and insects are shaped by sexual selection. Males acoustically compete for territories, and females choose their mates by means of male courtship songs. In courtship, richness and complexity of elements are often favoured characters. Only a few examples of complex songs are known in mammals. Males of the harem-polygynous sac-winged bat (Saccopteryx bilineata, Emballonuridae) have an uncommonly complex vocal repertoire, and different song types of males are used in the context of territorial defence and in courting females. We classified the daytime vocalisations of 16 male S. bilineata from a colony in Costa Rica, both on the basis of their acoustical properties and the social context in which they occurred. Seven vocalisation types were differentiated: echolocation pulses, barks, chatter, whistles, screeches, territorial songs and courtship songs. Territorial songs were short, rather stereotyped and not obviously directed towards a certain conspecific. They appear to be of importance in male competition for harem territories, in which females roost during the day. Courtship songs were exclusively observed when males displayed towards a female; they were long and complex, and consisted of highly variable elements (calls). We classified the calls in courtship songs of six males into call types, based on acoustical properties, mainly spectral purity and duration. Four call types are described in detail: trills, noise-bursts, short tonal calls, and quasi constant frequency calls. Twelve parameter values were extracted from the most common call type, the trill. Discriminant function analysis of trills showed that different males had different repertoires. This could allow females to use trill parameters for recognition of individual males and thus for mate choice.Communicated by G. Wilkinson     

Age-specific social dominance affects habitat use by breeding American redstarts (Setophaga ruticilla): a removal experiment

Summary By removing older males from their breeding territories, we tested the hypothesis that age-related dominance behavior influenced the pattern of habitat selection by breeding American redstarts Setophaga ruticilla (Aves: Parulinae). Fifteen older male redstarts removed in five experimental replicates during three breeding seasons were replaced by ten yearling and five older males; no redstart males of either age colonized the control sites during the same time periods, although two yearlings disappeared. Significantly more yearling males (67%, n=9) colonized the vacated areas than were present in the redstart population at large (26.8%, n=209). We reject the alternative hypothesis that yearling male redstarts occupy different habitats from older males because of age-related (innate) habitat preferences. Redstarts that colonized the territories made vacant by our removals (i.e., floaters) were a behaviorally heterogeneous group of animals. The presence of both yearling and older male floaters indicates that suitable habitat is limiting for this species and that intraspecific competitive interactions are important in habitat distribution, and potentially in population regulation.     

Compositional topography of melon and spermaceti organ lipids in the pygmy sperm whale Kogia breviceps: Implications for echolocation

The forehead of the pygmy sperm whale Kogia breviceps contains a large melon of fatty tissue in front of a small, fat-filled, cornucopia-shaped spermaceti organ. This unique anatomical structure may possibly play an acoustical role in the whale's echolocation system, similar to the fatty melon sound lens postulated for dolphins. To better understand its function, we have studied the compositional topography of the K. breviceps melon and spermaceti organ lipids. The fatty head tissues of an adult K. breviceps were serially sectioned into 9 transverse slices. Appropriate tissue samples were cut from every other slice, and analyzed for % lipid and lipid class composition. Wax esters and triglycerides were the only major lipids present; their average carbon number in each sample was determined by gas-liquid chromatography (GLC). Our topographical analyses of K. breviceps melon indicate 3 regions of distinctive lipid composition: a fat-poor melon exterior, an outer melon of medium fat content having % triglyceride>% wax ester, and a fat-rich inner melon having % wax ester>% triclyceride. The spermaceti organ contains a fat-rich core of very high wax-ester content (84 to 99%), surrounded by a fat-poor case. Average carbon numbers of both wax esters and triglycerides were lowest in the inner melon and the spermaceti organ. At the rear of the spermaceti organ lies the museau de singe, an apparent sound generator. The lipid topography data plus anatomical and acoustical considerations suggest that the K. breviceps melon/spermaceti organ system may function as an acoustical transducer, directing and refracting sound waves from this source for the purpose of echolocation.     

Risk-taking by female ducks: intra- and interspecific tests of nest defense theory

We tested several predictions of nest defense theory by observing variation in flushing distance and probability of nest abandonment within and between six species of waterfowl. In these species, only the females incubate eggs and attend offspring. First, we examined whether flushing distance by females varied in relation to clutch size, stage of incubation, and time of season, after controlling for the number of visits made to nests by observers. Revisits by observers appeared to affect flushing distance by females for reasons unrelated to the relative value of the current clutch. We found that as incubation progressed, females allowed observers to approach more closely before flushing from the nest. In some species, females with larger clutches allowed closer approaches to nests before flushing which was also consistent with nest defense theory. In contrast, time of season (Julian date) did not relate to flushing distance for any species. When species were compared, we found that species with moderate to high yearly mortality and high reproductive output per breeding attempt (e.g., northern shoveler and blue-winged teal) were less likely to abandon nesting attempts and exhibited riskier behavior (remained at nests when approached closely by observers) than species that had lower yearly mortality (e.g., mallard). Our results show that flushing distance and patterns of nest abandonment by female ducks conform to several predictions of nest defense theory.Correspondence to: M.R.L. Forbes     

Experimental studies on water strider mating dynamics: spatial variation in density and sex ratio

Summary We used field surveys, field experiments and experiments in artificial pools to study the effects of variation in sex ratio and density on mating dynamics of a stream water strider, Aquarius remigis. Our field survey documented the existence of hot spots, sites of higher than average total gerrid density, a male-biased sex ratio, and higher than average female mating activity. Female gerrids frequently changed sites, usually moving upstream, perhaps to spread their eggs among many sites. Male gerrids showed two movement strategies: some males frequently changed sites, while other males were stationary at hot spots. Surprisingly, smaller males tended to be stationary at hot spots. A field manipulation of the availability of refuges for females to avoid harassment by males supported the notion that males prefer hot spots because they are sites where a scarcity of refuge for females makes it relatively easy for males to intercept females. Experiments in plastic pools compared sites with 20 males: 5 females (simulating hot spots) to pools with 5 males: 5 females. The rate of male harassment of females was higher in 20:5 pools as compared to 5:5 pools. In response to increased male harassment, females reduced their activity on the water and increased their time spent out of the water and thus unable to forage. Males showed a large male mating advantage (LMMA) in 5:5 pools, but, surprisingly, not in 20: 5 pools. This pattern can explain the field observation that small males prefer hot spots. A behavioral mechanism that can explain the LMMA is as follows. Mating occurs when males overcome female resistance. Larger males have a mating advantage over smaller males if females resist heavily. Increased harassment (e.g., in 20:5 pools as compared to 5:5 pools) might result in reduced female resistance to males and thus a reduced LMMA. Females also showed some non-random mating by size that might reflect an interplay between male preference for large females and the avoidance of males by large females. Correspondence to: A. Sih     

Primary productivity and sizes of pools of organic carbon in the mixed layer of the ocean

We estimated primary productivity and distributions of carbon in the phytoplankton, micro-zooplankton, and suspended and dissolved matter in various areas of the World Ocean to increase our information about the organic carbon cycle in the surface layer of the sea. Primary productivity ranged from about 0.1 gC m^–2 day^–1 in the Gulf of Mexico to 9 gC m^–2 day^–1 in nutrient-rich water off Peru. Phytoplankton carbon ranged from less than 10 g/l in the former to 750 g/l in the latter and in nutrient-rich water off southwest Africa. Micro-zooplankton carbon usually was less than 50 g/l in all waters, and was dominated by ciliates, copepodids, and copepod nauplii in all areas. Concentrations of particulate carbon ranged from 12 g/l off the east coast of South America to 850 g/l off southwest Africa. Concentrations of dissolved organic carbon varied between 0.5 and 1.5 mg/l in all areas except off Peru, where maximum values of 4.5 mg/l were observed. Turnover rates of carbon by small standing crops of micro-flagellates (1 to 5 longest dimension) and dinoflagellates in nutrient-poor waters were lower than those by large standing crops of diatoms and micro-flagellates in nutrient-rich waters. Concentrations of phytoplankton usually accounted for 20 to 55% and micro-zooplankton for 2 to 30% of the particulate carbon in the surface layer of the sea. Concentrations of dissolved organic carbon were not related to concentrations of particulate carbon in most waters except off Peru, where they appear to be directly related.     

Multiple sex-chromosome system and other karyological characterizations of Pterotrachea hippocampus (Mollusca: Mesogastropoda)

Two modal diploid numbers of chromosomes were found for Pterotraches hippocampus Philippi (Mollusca: Mesogastropoda) collected from the Gulf of Palermo in 1990: 2n=31 and 32 for males and females, respectively. This, along with other karyological characteristics such as the occurrence of a trivalent configuration at diakinesis and two types of metaphase-II spreads in spermatocytes, supports the notion that a X₁X₂Y/X₁X₁X₂X₂ sex mechanism operates in the species investigated here. Silver nitrate procedure revealed an intraindividual variation in the Ag-staining pattern occurring in this species. The majority of the chromosome pair displayed terminal and/or interstitial heterochromatic blocks after C-banding.     

Residues of Organochlorinated Pesticides in Eggs of Water Birds from Tai Lake in China

The levels of organochlorine compounds in eggs of water birds from the colony on Tai Lake in China were studied. The eggs were collected in 2000 and belonged to the following species: 65 samples of black-crowned night heron (Nycticorax nycticorax), 36 samples of little egret (Egretta garzetta), 26 samples of cattle egret (Bubulcus ibis) from 13 clutches and 43 samples of Chinese pond heron (Ardeola bacchus) from 17 clutches. Dichlorodiphenyltrichloroethane (DDT) and its derivates (DDE and DDD), hexachlorocyclohexane (HCH) and its isomers (-HCH, -HCH, -HCH, -HCH), heptachlor, heptachlor epoxide, aldrin, dieldrin, endrin, endrin aldehyde, -endosulfan, -endosulfan, and endosulfan sulfate were determined in the laboratory by gas chromatography. The data showed that DDE had the highest levels in all the samples, followed by -HCH. The mean levels of DDE among the water bird species were in the order as follows: black-crowned night heron (5464.26ng/g, dry weight) > Chinese pond heron (2791.12ng/g, dry weight) > little egret (1979.97ng/g, dry weight) > cattle egret (660.11ng/g, dry weight). DDT and its metabolites accounted for 90% of the total organochlorines, except that it was only 73% for cattle egret. The differences of the residue among the bird species were statistically significant and could be attributed to their variations in prey and habitat. Although the DDE burdens in Tai Lake were much lower than 8 g/g (wet weight) which are thought to have significant adverse effects on black-crowned night herons, they would be expected to increase the risk of adverse effects on survival of chicks of herons and egrets, particularly black-crowned night heron, based on the critical value of 1 g/g (wet weight) DDE. The burdens of HCHs in this study were higher and the cyclodienes were lower than those found elsewhere.     

Pillar building in the fiddler crab Uca beebei: evidence for a condition-dependent ornament

In the fiddler crab (Uca beebei) males build a small mud pillar next to their burrow which increases their attractiveness to females. Three hypotheses were tested to explain inter-male variation in pillar-building. (1) The benefits of pillar-building are density dependent. The experimental addition of vertical structures did not support this hypothesis as there was no change in the level of pillar-building. (2) There are two classes of males (pillar-builders and non-pillar-builders). This could either be due to an alternative mating strategy, or because pillar building is age or size-dependent. There was also no support for this hypothesis. (3) Pillar-building is an honest signal of male quality dependent on body condition. A food supplementation experiment was performed. Addition of food affected several aspects of male behaviour and resulted in a two fold increase in the number of pillars built between control and food treatments (P < 0.001). However, the percentage of males building pillars did not increase significantly. Pillar building in this species has been attributed to sensory exploitation. Our results indicate that a trait which may well have evolved through sensory exploitation also appears to be condition-dependent. We emphasise that showing that an ornament or behaviour is condition-dependent does not necessarily mean that it evolved through good gene processes. However, in terms of its current selective value, pillar building may be maintained through female choice because it acts as a signal of male condition.