Delayed breeding in the cooperatively breeding Florida scrub-jay (Aphelocoma coerulescens): inhibition or the absence of stimulation?

To determine whether fundamental differences exist in the reproductive physiology of breeder and nonbreeder Florida scrub-jays (Aphelocoma coerulescens), we compared plasma levels of testosterone (T) and luteinizing hormone (LH) in males, and estradiol (E₂) and LH in females. Although male breeders had higher overall T and larger testes, nonbreeders’ T paralleled that of breeders, and their testes were more than an order of magnitude larger than regressed testes. Breeder and nonbreeder males had equivalent baseline LH, and equivalent changes in LH following a gonadotropin-releasing hormone (cGnRH-I) challenge. The T, LH and GnRH challenge data indicate that nonbreeder males have functional hypothalamo-pituitary-gonadal (HPG) axes. We found no hormonal evidence of inbreeding suppression in males: nonbreeders that did not live with their mothers and those that did had similar T. Male nonbreeders that were exposed to E₂-implanted females had higher T than did controls, suggesting that the lack of within-pair stimulation is a key factor in whether an individual delays breeding. Female nonbreeders had E₂ titres equal to or higher than breeders and neither basal LH nor LH following GnRH challenge differed by breeding status. Nonbreeders’ ovarian follicles were smaller than breeders’, but were larger than they would be during the non-breeding season. These data suggest that nonbreeders were primed for breeding and were simply waiting for an opportunity or a required stimulus. Female nonbreeders that lived in a territory with an unrelated male breeder had significantly higher E₂ than those that remained with their fathers. Similarly, nonbreeders that were captured away from their home territories had elevated E₂. However, nonbreeders that lived with their fathers had E₂ that was equivalent to breeding females, suggesting that inbreeding avoidance may not be the primary factor leading to delayed breeding in females. Received: 13 June 1995 /Accepted after revision: 27 April 1996     

High paternal investment in unrelated young: extra-pair paternity and male parental care in house martins

The response of males to reduced paternity has important consequences for the evolution and maintenance of a mixed reproductive strategy. Paternity is predicted to affect directly the level of male parental care in some cases but not in others. The response of males to reduced paternity will be influenced by their ability to assess their paternity, the predictability of cuckoldry and the costs and benefits of parental care. Although male house martins (Delichon urbica) provide among the highest levels of male parental care known in passerines (incubation, brooding and feeding nestlings), there was no evidence that cuckolded males substantially reduced their level of parental care, and, as a result, all young fledged successfully. Thus, extra-pair fertilizations enhanced the reproductive success of some males because they were able to parasitize the parental care of cuckolded males. We discuss several conditions which may favor extensive male parental care even when the male's paternity is very low.     

Mating effort or paternal investment? Incorporation rate and cost of male donations in the wartbiter

Summary Provision of nuptial gifts and incorporation of male-derived substances occurs in several insects. There has been some discussion as to whether these gifts should be regarded as a form of paternal investment or part of mating effort. One assumption of the paternal investment hypothesis is that the donating male is expected to father all, or the majority of, the offspring. The rate of incorporation into developing eggs is therefore assumed to be rapid enough to assure the donating male of paternity of the offspring in which he is investing. Spermatophore production in a paternal investment system is also expected to be costly, since males are providing females with highly nutritious gifts during copulation, thereby increasing female fecundity. Therefore, under conditions of food limitation males are expected to alter either mating frequency or spermatophore size. Experiments on the wartbiter (Decticus verrucivorus), using radioactive isotopes, show that the incorporation rate of male-derived substances is so slow that females become sexually receptive before any material received from the last male have been incorporated into eggs laid by the female. Since sperm mixing occurs in this species and since the label is present in eggs laid up to more than 30 days after mating, males might potentially benefit another male's offspring. The nuptial gift (the spermatophylax) was found, regardless of mating frequency and diet, to have a low protein content. These findings coincide with previous results showing that consumption of the gift had no effect on female fecundity. Instead, wartbiter males keep mating frequency high, are capable of mating every day regardless of food availability, invest the same proportion of the body weight in spermatophore production regardless of size, and produce a spermatophylax that correlates with ampulla size. These results indicate that the nuptial gift mainly functions to increase fertilization success in the wartbiter.     

Maternal investment in rhesus macaques (Macaca mulatta): reproductive costs and consequences of raising sons

Maternal investment in offspring is expected to vary according to offspring sex when the reproductive success of the progeny is a function of differential levels of parental expenditure. We conducted a longitudinal investigation of rhesus macaques to determine whether variation in male progeny production, measured with both DNA fingerprinting and short tandem repeat marker typing, could be traced back to patterns of maternal investment. Males weigh significantly more than females at birth, despite an absence of sex differences in gestation length. Size dimorphism increases during infancy, with maternal rank associated with son’s, but not daughter’s, weight at the end of the period of maternal investment. Son’s, but not daughter’s, weight at 1 year of age is significantly correlated with adult weight, and male, but not female, weight accounts for a portion of the variance in reproductive success. Variance in annual offspring output was three- to fourfold higher in males than in females. We suggest that energetic costs of rearing sons could be buffered by fetal delivery of testosterone to the mother, which is aromatized to estrogen and fosters fat accumulation during gestation. We conclude that maternal investment is only slightly greater in sons than in daughters, with mothers endowing sons with extra resources because son, but not daughter, mass has ramifications for offspring sirehood. However, male reproductive tactics supersede maternal investment patterns as fundamental regulators of male fitness. Received: 23 July 1999 / Received in revised form: 23 February 2000 / Accepted: 13 March 2000     

Toward a delineation of the circumstances in which cooperation can be sustained in environmental and resource problems

Play in standard laboratory Public Good games suggests that on average, humans are quite prone to cooperate. Yet cooperation is often absent in real world social dilemmas, including many environmental problems. We propose that this discrepancy arises because in the Public Good game, the worst freeriders can do is to not contribute to the public account, while in many real world environmental situations freeriders can even appropriate contributions made by others before the public good is produced. We introduce the Claim Game that modifies the Public Good game by allowing for appropriating the contributions of others before the public good is produced. The impact of such possible takings on public good production is dramatic. No public good is produced, not even in the initial stages of interaction. We link our findings to the relevance of common pool games for modeling environmental problems, and stress the need to experimentally test environmental institutions within harsher social dilemmas than the standard Public Good game.     

Traditional management of lagoons for fishery can be inconsistent with restoration purposes: the Valli di Comacchio study case

We gathered water and macrobenthic data over a 5-year period in the Valli di Comacchio, the largest lagoons in the Po River Delta (northern Adriatic), to evaluate the response of ecological indices and to test their sensitivity against environmental variables. Of 84 sampling points, only 12 exhibited a good/high ecological quality status according to the BITS and 28 according to the M-AMBI. Environmental variables representative of eutrophication did not correlate with the indices. The impacts experienced by the Valli during the last 30 years seem to have led to disruption of the pelagic–benthic food web. Eel fisheries, whose management is traditionally characterised by long water residence time, thus seem inconsistent with restoration purposes.     

Sexual dimorphism in the buccal cavity of paternal mouthbrooding cardinalfishes (Pisces: Apogonidae)

The mechanical basis of mouthbrooding in the coral reef cardinalfish (family Apogonidae) has yet to be established. Sexual dimorphism in the anatomy of the oral jaws and buccal cavity of seven species of paternal mouthbrooding apogonids was therefore investigated. A novel silicon injecting technique was used to quantify between sexes differences in size and shape of the buccal cavity. Osteological and morphological measurements were also examined to identify the structural features associated with variation in buccal volumes. Males buccal volumes were proportionally larger by 20–42% in five of the seven species (P<0.01), a change that was largely a result of an increase in the depth of buccal cavity. Absence of distinct osteological differences between sexes suggests that changes in the soft anatomy are primarily responsible for the increased buccal volume in males. The average increase of 31% in buccal volume of males appears to reflect their role in mouthbrooding. This is the first study to demonstrate sexual dimorphism of the buccal cavity of multiple mouthbrooding species.     

Deviations from predictions of the metabolic theory of ecology can be explained by violations of assumptions

The metabolic theory of ecology (MTE) is based on models derived from the first principles of thermodynamics and biochemical kinetics. The MTE predicts that the relationship between temperature and species richness of ectotherms should show a specific slope. Testing the validity of this model, however, depends on whether empirical data do not violate assumptions and are obtained within contour conditions. When dealing with richness gradients, the MTE must be empirically tested only for ectothermic organisms at high organization levels and when their body size as well as abundance does not vary with temperature gradients. Here we evaluate whether the magnitude of the deviations in slope expected from the MTE to empirical data for New World amphibians is due to the violations of model assumptions and to lack of generality due to restricting contour conditions. We found that the MTE correctly predicted biodiversity patterns only at higher levels of organization and when assumptions of the basic model were not violated. Approximately 60% of the deviations from the MTE-predicted slope across amphibian families were due to violations of the model assumptions. The hypothesis that richness patterns are a function of environmental temperature is too restrictive and does not take complex environmental and ecological processes into account. However, our results suggest that it may be possible to obtain multiple derivations of the MTE equation if idiosyncrasies in spatial and biological/ecological issues that are essential to understanding biodiversity patterns are considered.     

Targeting Ecosystem Features for Conservation: Standing Crops in the Florida Everglades

Abstract: The Everglades in southern Florida, U.S.A., is a major focus of conservation activities. The freshwater wetlands of the Everglades do not have high species richness, and no species of threatened aquatic animals or plants live there. We have, however, identified a distinctive ecological feature of the Everglades that is threatened by canal construction, draining, and nutrient enrichment from agricultural runoff. Compared to values reported from other freshwater systems, standing stocks of periphyton in relatively undisturbed areas of the Everglades were unusually high, and standing stocks of invertebrates and fish were unusually low. Averaging data gathered from nine sites and five sampling periods spanning 1 year, we found that periphyton standing crop was 88.2 g/m² (ash-free dry mass), invertebrate standing stock was 0.64 g/m² (dry mass), and fish standing stock was 1.2 g/m² (dry mass of large and small species combined). We found that fish standing stocks were much higher in phosphorus-enriched sites than in nearby reference sites but that invertebrate standing stocks were similar in enriched and reference sites. Our results support the notion that oligotrophy is at least partially responsible for the low standing stocks of fish, but they also suggest that species interactions and a paucity of deep-water refugia are important. Anthropogenic eutrophication in Everglades marshes will lead to the loss of distinctive ecosystem features. A focus on species richness and "hot spots" of threatened species provides no basis for conservation of ecosystems like the Everglades. If oligotrophic ecosystems often have low species richness, they will be underrepresented in preservation networks based on some common criteria for establishing conservation priorities.     

Parentage and paternal care: consequences of intersexual selection in Savannah sparrows?

Empirical relationships between parentage and male parental care are commonly interpreted in the context of life-history models that consider increased offspring survivorship as the only benefit of paternal effort. However, indirect benefits associated with male care can also influence a male's response to cuckoldry: if females allocate paternity according to their prior experience with male parental care, it may pay for males to provision extra-pair young in early broods. Here, I assess the relationship between first-brood parentage and paternal care in a population of Savannah sparrows (Passerculussandwichensis) where a male's fertilization success in the second brood appears to be influenced by his prior parental performance. Based on the multi-locus DNA fingerprinting of 17 first broods, male feeding effort was influenced by parentage (percent of brood resulting from within-pair fertilizations) but not by brood size, male mating status (monogamous versus polygynous), timing of breeding (hatching date), structural size (wing length) or condition (mass). Males provided more care to broods that contained few within-pair young. This result supports the idea that males provision young to increase their future mating success, but alternative hypotheses involving male quality and timing of breeding cannot be excluded. Received: 13 August 1996 / Accepted after revision: 22 February 1997     

Time and recruitment costs as currencies in manipulation studies on the costs of reproduction

Life history theory predicts that parents will have lower Darwinian fitness if they tend clutches that are above or below the size they naturally produce. We experimentally tested for relationships between fitness and clutch size in Tree Swallow (Tachycineta bicolor) offspring and parents. Over 130 trios of nests initiated on the same day were randomly divided among reduce (-3 eggs), control (3 eggs picked up and replaced), or add (+3 eggs) manipulations. Pre-manipulation modal clutch size was six eggs (range before manipulations was 1-10; afterwards, it was 1-11). Hatching took longer in larger clutches, but the proportion of eggs hatching and fledging was similar for clutches from 4 to 10, so that clutches of 10 produced the maximum number of fledgling. Parental feeding rates were higher for larger broods, but per capita feeds to nestlings were fewer, and nestlings were smaller. Nonetheless, survival of both young and adults, based on recaptures in subsequent years, was not significantly affected by manipulations. Manipulations also had no significant effect on subsequent reproduction, including the number of fledglings produced by either local recruits or returning breeders. Collectively, our results failed to detect fitness costs associated with tending larger clutches for either parents or the offspring reared and suggested directional selection for larger clutch size. However, because clutches that hatch later produce fewer recruits, the extra days required to lay more eggs and to fledge extra young may eliminate a large part of the advantage that would accrue to parents producing enlarged clutches. For example, our data suggest that there may be less than a 16% benefit to producing nine instead of six eggs, rather than 50%, as is suggested by experimentally manipulated egg numbers alone. Thus, time, rather than costs of reproduction, may be the crucial constraint selecting against Tree Swallows laying larger clutches.     

Life and Death in the Fast Lane: Demographic Consequences of Road Mortality in the Florida Scrub-Jay

Abstract: We examined the demographic consequences of road mortality in the cooperatively breeding Florida Scrub-Jay (Aphelocoma coerulescens ), a threatened species restricted to the oak scrub of peninsular Florida. Between May 1986 and July 1995 we monitored the survival and reproductive success of a color-banded population of jays along a two-lane highway at Archbold Biological Station. Annual mortality of breeding adults was 0.38 on road territories, significantly higher than the rate of 0.23 for breeders on nonroad territories. High mortality on road territories appeared to be a direct result of automobile traffic per se and not a consequence of road-induced changes in habitat characteristics. Mortality was especially high for immigrants without previous experience living along the road: in their first two years as breeders on road territories, naive immigrants experienced annual mortality of 0.50 and 0.45. From year 3 onward, however, annual mortality dropped to 0.29, not significantly different from the rate for birds on nonroad territories. This experience-dependent decline in road mortality could be caused either by surviving jays learning to avoid automobiles or by selective mortality operating through time (demographic heterogeneity). Proximity to the road had no effect on nesting success beyond its indirect effects on breeder experience and group size. Because the mortality of 30- to 90-day-old fledglings was significantly higher on road territories than on nonroad territories, however, breeder mortality greatly exceeded production of yearlings on road territories. Roadside territories therefore are sinks that can maintain populations of Florida Scrub-Jays only via immigration. Because Florida Scrub-Jays do not avoid roadside habitats and may even be attracted to them, road mortality presents a difficult challenge for the management and conservation of this threatened and declining species.     

Evidence that lipid can be digested by the dumpling squid Euprymna tasmanica, but is not stored in the digestive gland

The aims of this study were to assess quantitatively the enzymatic ability of squid to digest lipids and the ability of the digestive gland to accumulate lipid classes associated with storage. This was achieved through two manipulative experiments using the dumpling squid, Euprymna tasmanica. Firstly, we measured lipase activity and determined the presence and location of lipid vacuoles within the digestive gland; secondly we identified and quantified lipid classes in the digestive gland. Given the levels of lipase activity, we provided evidence for the first time that a squid species is capable of digesting lipid at levels comparable to invertebrates known to use dietary lipid. A poor relationship between feeding activity and lipase secretion suggests that enzyme production is continuous. The second experiment found no evidence that lipid was stored in the digestive gland; most of the lipid present in the gland was either structural or a dietary by-product. The implication of these findings is that for this species lipid is most probably being immediately digested and used for growth and reproduction rather than being stored in the digestive gland. We consider that the role and storage of lipid is likely to vary among different cephalopod species, but not predictably as function of their lifestyle. Therefore, potential locations for lipid storage, other than the digestive gland, need to be considered and using changes in the relative size of the digestive gland as a measure of condition needs to be interpreted with care.     

Maternal investment in sons and daughters: problems of methodology

Summary The prediction that mothers should invest more in sons than daughters was briefly examined in American bison (Bison bison) by Wolff (1988). He concluded that (a) male calves suckle more than females, (b) cows that had previously produced sons were more likely to be barren, and (c) cows that had male calves became estrous later in the year than other cows. In this paper we present data from two long term studies at different sites to show that Wolff's conclusions are equivocal at best and difficult to reconcile with predictions of the hypothesis because of questionable methods and assumptions. Offprint requests to: W.C.H. Green     

Mixed-species shoaling in fish: the sensory mechanisms and costs of shoal choice

The mechanisms and functions of mixed-species shoaling were investigated in two sympatric species of cyprinids, the chub and the European minnow, from the river Wharfe where they comprised approximately 70% of all year 0+ fish over a 20-year survey. Chub preferred conspecific shoals over heterospecific ones with olfactory cues being more important than visual ones for shoal choice. This preference was consistent with measurements of length:flank area ratios and length:weight ratios which suggest that both species are similar in appearance. When presented with mixed-species shoals, chub increased the percentage time spent with stimulus shoals with increasing proportions of conspecifics. Feeding experiments suggest that the preference for conspecific shoals is driven by interspecific competition (with minnows out-competing similar-sized chub) and the oddity effect. The importance of this work in the context of species assortment in free-ranging shoals is discussed.     

Rarity of Organisms in the Sand Pine Scrub Habitat of Florida

Naturalists conceive of "rare" organisms as those narrow in geographic range, restricted to few habitats, or small in population sizes. Because rare organisms may be rare in a variety of ways, a particular conservation strategy that purports to protect them may not do so effectively in all cases; therefore, categorization of rarity in advance could help ensure that the chosen strategy has a reasonable chance of success in a particular case. We placed plants, amphibians, and reptiles resident in the Sand Pine scrub habitat of Florida into categories of rarity. Determining if and in what way a particular organism is rare proved difficult, because the process requires exact specification of what is meant by "narrow,""restricted," and "small." Furthermore, the process requires choice of a scale for judging rarity and relies upon data that are likely to be of variable availability and reliability. We categorized rarity by several schemes, and they produced distributions of taxa among categories of rarity that largely were similar to one another but that diverged in telling ways. The distributions of taxa among categories of rarity produced by any scheme were different for scrub plants than for scrub amphibians and reptiles. Likewise, these distributions were different than those produced by previously published studies of rarity. We found that the results of our categorization schemes did not match particularly well with the official listing of scrub organisms as endangered or potentially endangered.     

Optimal pollution in the presence of adjustment costs

This paper explores optimal pollution control when pollution is considered a stock and costs of treating pollution are a function both of the level and rate of change of pollution control. It is shown that even if the long run optimum stock of pollution is below the initial level adjustment costs may imply an optimally increasing stock of pollution in the short run. The results obtained are contrasted with those which emerge when adjustment costs are absent and when damages are taken to be a function of the flow of pollution; the appropriate tax rates of polluting discharges are examined.     

Orchid bees don't need orchids: evidence from the naturalization of an orchid bee in Florida

Almost 200 species of orchid bees are the exclusive pollinators of nearly 700 specialized orchids in the neotropics. This well-known mutualism involves orchids, called perfume orchids, which produce species-specific blends of floral fragrances, and male orchid bees, which collect and use these fragrance compounds during their courtship. We report here the naturalization of an orchid bee, Euglossa viridissima, in southern Florida, USA, where perfume orchids are absent. Chemical analysis of the contents of the fragrance storage organs in the hind tibias of 59 male bees collected in Florida identified 55 fragrance compounds, including 27 known from the perfumes of nine species of E. viridissima's orchid mutualists in Mesoamerica. Aromatic leaves, such as basil, were found to be important surrogate sources of needed fragrance compounds in Florida. The bee's ability to live and become abundant in the absence of its orchid mutualists suggests that the orchid bee-perfume orchid mutualism may be facultative for the bees, even though it is obligatory for the orchids. This invasive bee visits and potentially pollinates the flowers of many plants in Florida, behavior that could promote the abundance of selected exotic and native species.     

Maternal investment in Antarctic fur seals: evidence for equality in the sexes?

Studies of the otariids (fur seals and sea lions), a highly sexually dimorphic group, have provided conflicting evidence of differential maternal expenditure in male and female offspring and, thus, suggestions that they conform to predictions of investment theory are equivocal. Since the mid-1970s, a diversity of research on Antarctic fur seals (Arctocephalus gazella) including studies of their reproductive ecology, lactation energetics, and foraging behaviour have been conducted at Bird Island, South Georgia that have resulted in one of the more complete and diverse data sets for any species of otariid. These long-term data were reviewed to determine whether there was any evidence to support that differential maternal expenditure occurred in Antarctic fur seals. Most of the data examined were collected during five consecutive austral summers from 1988 through 1992 and included years in which local food resources were abundant and scarce. We were unable to detect differences in the sex ratios of pups at birth or sex-biased differences in growth rates estimated from serial data, the number of foraging trips made, the duration of attendance ashore, diving behaviour, suckling behaviour, or milk consumption in any year and in the duration of foraging trips or age at weaning in 2 of 3 years. In addition, we found no evidence of greater reproductive costs between mothers with sons or daughters relative to their reproductive performance the following year. In contrast, sex-biased differences were only found in the duration of foraging trips in 1990, the age at weaning in 1988, and consistently in growth rates estimated from cross-sectional data. We suggest that differential maternal expenditure does not occur in Antarctic fur seals because male pups probably do not gain greater benefit from additional maternal expenditure than female pups. After weaning, males experience a period of rapid juvenile growth over 3–4 years during which time body mass nearly trebles. This growth will almost certainly be dependent upon available food resources then rather than on any maternal expenditure received over the first 4 months of life and, thus, the assumptions of the Trivers and Willard hypothesis are probably invalid for Antarctic fur seals. Received: 10 July 1996 / Accepted after revision: 3 March 1997