Linking micromorphism,brooding, and hermaphroditism in brachiopods: Insights from Caribbean Argyrotheca (Brachiopoda)

In extant brachiopods, parental brooding of the larvae occurs exclusively within Rhynchonelliformea. Methods of larval protection range from simple retention of the larvae within the mantle cavity, to sophisticated brood care within highly specialized brood pouches found in Argyrotheca and Joania (Terebratulida, Megathyridoidea), Gwynia (Terebratulida, Gwynioidea), and all Thecideoidea (Thecideida). Previous studies on the reproductive biology of Argyrotheca yielded contrasting results on the epithelial origin of the brood pouches in this genus. Here, representatives of different species of Argyrotheca from the Belize Barrier Reef were examined using histological section series. Brood pouches of four species, A. cf. schrammi and Argyrotheca sp. 1–3, are of the same basic structure, formed by invaginations of the anterior body wall and connected to the visceral cavity via the metanephridia. The same four species are simultaneously hermaphroditic, suggesting that fertilization is achieved, at least partly, through selfing. One species, Argyrotheca rubrocostata, differs significantly from all others as it has no brood pouch and gonochoric gonads. Thus, the presence of brood pouches and simultaneous hermaphroditism are concluded to be correlated within Megathyridoidea and proposed to be homologous traits of Joania and several but not all species of Argyrotheca, questioning the monophyletic status of both genera. In contrast to the brood pouches of Thecideoidea, lophophoral epithelium is not involved in the formation of the pouches of Argyrotheca and Joania. Therefore, megathyridoid and thecideoid brood pouches are not homologous but evolved independently within rhynchonelliform brachiopods. All brachiopods with brood pouches share a micromorphic form and a short life span, limiting the space and time available for gamete and larval development. We suggest that the brood pouches and the hermaphroditic gonads of Argyrotheca spp. and Joania compensate these limitations by minimizing the loss of gametes and larvae, and by maximizing the chances of successful fertilization. J. Morphol., 2013. © 2013 Wiley Periodicals, Inc.     

Sociality in the phylogenetically basal allodapine bee genus Macrogalea (Apidae: Xylocopinae): implications for social evolution in the tribe Allodapini

Recent phylogenetic studies based on DNA sequence data indicate that the tropical African bee genus Macrogalea is the sister group to the remaining extant allodapine fauna, whereas previously it was thought to be a distal group. This leads to some fundamental changes in our understanding of social evolution in the allodapines. Earlier studies suggested that Macrogalea showed only weak forms of social behaviour and these were not well characterized. However, large samples of Macrogalea zanzibarica presented here show that this species exhibits marked social behaviour. Nearly half of nests collected contained two or more adult females, with up to 10 females per nest. Brood are reared progressively and brood ages within colonies are staggered, giving rise to colonies with very mixed age structures and therefore frequent opportunities for alloparental care. Ovarian dissections indicate non‐simple forms of reproductive partitioning within colonies and most multi‐female colonies show evidence that more than one female has contributed to egg production, though reproductive episodes among colony members are frequently asynchronous. Some females show signs of much higher wing wear than their nestmates, but always show signs of previous reproduction. Reproductive division of labour appears to be temporally marked, ovarian differentiation among nestmates is linked to relative body size, but permanent worker‐like or queen‐like castes appear to be absent. This is similar to the communal, continuously brooded and multivoltine behaviour of some tropical halictine species and may be due to the aseasonal nature of brood development in tropical regions. Patterns of per capita brood production indicate large benefits to multi‐female nest occupancy, and sex allocation is strongly female biased. These findings strongly suggest that the capacity for complex social interactions and alloparental care is an ancestral trait for all of the extant allodapine lineages. Therefore comparisons among extant allodapines are unlikely to throw light on the initial origin of social behaviour, though they may uncover origins of true caste behaviour and reversals to solitary nesting. Sex ratios in Macrogalea and most other allodapine genera, spanning a broad phylogenetic and ecological range, suggest that one or more allodapine traits have provided persistent selection for female‐biased sex allocation.     

Small hosts infrequently disrupt laying by Brown-headed Cowbirds and Bronzed Cowbirds

ABSTRACT Brood parasites often must overcome host defenses that may include behaviors that serve other functions, such as deterrence of predators and nest attendance during laying and incubation. Host use by brood parasites may also be influenced by competitors in areas where more than one parasitic species occurs. We identified the degree to which behavior of potential hosts and potential competitors affected laying by Brown‐headed Cowbirds (Molothrus ater) and Bronzed Cowbirds (M. aeneus) at a site in south Texas where they co‐occur. We watched potential host nests during the presunrise period to record cowbird laying and document nest visitation, laying, cowbird‐host encounters, and nest attentiveness by hosts. Hosts were frequently at their nests when cowbirds laid eggs (83% of 121 watches among nests of five host species) and cowbirds regularly encountered hosts (43–74% and 40–77% of watches per species of host for Brown‐headed and Bronzed cowbirds, respectively). Host nest defense infrequently interfered with cowbird laying and cowbirds rarely interacted with one another during laying. Overall, 12% of the 42 cowbird laying attempts that elicited host nest defense failed, resulting in cowbird eggs either laid atop hosts as they sat in nests or laid outside the nest cup. We clearly documented that relatively small hosts can thwart parasitism by cowbirds. Thus, the potential for successful defense of nests should be considered when assessing the evolution of behaviors to deter the removal of host eggs by cowbirds and mechanisms leading to nest abandonment. Regarding the latter, the presence of a cowbird at a nest would be a poor indicator for parasitism as some laying attempts were thwarted and unparasitized broods were reared at those nests. Despite the potential for nest defense to affect host use by cowbirds, we did not detect an effect of nest defense. Because most host defense was ineffective, we examined hypotheses for the timing of cowbird laying and host nest attendance. Our analysis of time of day of laying by Brown‐headed Cowbirds at our site and data compiled from the literature suggests that laying time is best predicted by the time of civil twilight (first light) rather than sunrise.     

Regulation of brood size in a burying beetle,Nicrophorus tomentosus (Silphidae)

Regulation of brood size in a biparental burying beetle, Nicrophorus tomentosusWeber, was studied by providing pairs with one of two sizes of mouse carcasses in the laboratory. For a given carcass size, there was an inverse relationship between number and mass offspring in a brood. The requirement for regulation was that brood size was adjusted such that mean mass of individual larvae was constant for carcasses of different size. Brood size was regulated if parents were present but regulation did not occur if parents were removed prior to hatching of larvae. Pairs bred in quick succession on two carcasses raised fewer than the regulated number of young in the second reproductive attempt. Reasons for regulation of brood size in this genus are discussed.     

Growth and reproduction of the mosquitofish,Gambusia affinis,in relation to temperature and ration level: consequences for life history

The allocation of energy to growth and reproduction, in relation to temperature and food availability, was investigated in laboratory experiments with the mosquitofish,Gambusia affinis. At constant temperature of 20, 25 and 30°C and ad libitum feeding, specific growth rates increased with increasing temperature at 1.7, 3.1 and 3.4% dry mass day⁻¹, respectively. Growth rates in a cycling temperature regime (20–30°C, ) were faster than in a 25°C constant temperature. As temperature increased from 20 to 30°C, mean age at first reproduction decreased from 191 to 56 days and brood size and mass of offspring increased significantly. Interbrood interval was also temperature dependent; estimates at 25 and 30°C for females >1000 mg were 22.6 and 18.6 days, respectively. Interbrood interval could not be calculated at 20°C. Although fitness was highest at 30°C, females at 25°C invested a greater proportion of surplus energy (growth and reproduction) to reproduction (38%) than at 20 (17%) or 30°C (36%) during the 32-week study. Fish at cooler temperatures began reproduction at a smaller size. Where rations were controlled at low, medium, and ad libitum levels, somatic and gonadal growth increased with increasing temperatures and food availability. The proportion of energy invested in reproduction was highest at 25°C for each comparable ration level. Calculated energy budgets indicated that over the 10-week study, 17–22% of the food energy was invested in growth, 0–7% in reproduction, and 75–83% in respiration and excretory losses, depending on feeding and temperature conditions.     

Temporal variation in abundance of the egg predator Carcinonemertes epialti (Nemertea) and its effect on egg mortality of its host,the shore crab,Hemigrapsus oregonensis

An outbreak of the nemertean, Carcinonemertes epialti, was observed on Hemigrapsus oregonensis during October, 1982 at Campbell Cove, Bodega Harbor, California. Mean worm intensity (296 worms/crab) was the highest recorded for this nemertean egg predator on H. oregonensis. During the outbreak, male crabs were found to harbor more worms than both non-ovigerous and ovigerous females. Crab egg mortality was substantial; 83% of the ovigerous females experienced 75–100% brood mortality. The seasonal peak in worm abundance coincided with the seasonal low in crab reproduction at this locality. A method for estimating the impact of C. epialti on H. oregonensis natality was developed using crab size and fecundity, and worm prevalence and intensity. For a non-outbreak sampling period, a mean of 5.6% egg mortality was experienced by infested crabs for the period selected. Thus, brood mortality during the outbreak was much greater than that experienced at non-outbreak periods. Heavy fishing pressure on some commercially important crab species has been suggested as a possible factor inducing worm outbreaks and facilitating their continued persistence. These observations suggest that fisheries are not necessarily responsible for the outbreaks of nemerteans on commercially important crab species. However, fishing pressure may still be a sufficient condition to promote nemertean outbreaks.     

Oxygen demand and long term changes of profundal zoobenthos

The paper attempts to combine the low oxygen content of the hypolimnion during stratification and the oxygen uptake of zoobenthos. Data of declining oxygen content in the hypolimnion and critical limits of respiration are combined for Chironomus anthracinus, Potamothrix hammoniensis and three species of Pisidium, P. casertanum, P. subtruncatum and P. henslowanum. The respiratory adaptation to low oxygen content influences both growth and population dynamics of the different species. The results have important bearing on eutrophication of the Lake Esrom ecosystem and temperate eutrophic lakes in general as well as the composition of profundal zoobenthos and its population dynamics.Publication No. 389 from Freshwater-Biological Laboratory, University of Copenhagen.     

Dominance in demand-feeding behaviour in Arctic charr and rainbow trout: the effect of stocking density

The effects of stocking density on the demand feeding behaviour of Arctic charr Salvelinus alpinus and rainbow trout Oncorhynchus mykiss were evaluated using demand feeders in combination with a monitoring system based on automatic passive integrated transponders. The proportion of total bites accounted for by top-ranked charr and trout decreased from 87 to 15% and from 66 to 15%, respectively, when moving from the lowest to the highest densities, indicating that the ability of the highest-ranked fish to monopolize the demand feeders was reduced at high densities. However, when the individuals were grouped into quartiles, based on their individual rank, both charr and trout ranked within the upper quartile accounted for the majority of bites independent of stocking density (87 and 72% of the total, respectively). Instead of there being a single dominant individual, as was the case under low density conditions, a small group of individuals dominated the actuation of the trigger at high densities. The two species responded differently to crowding, with the total daily bite activity of charr increasing linearly with increasing density, whereas the corresponding relationship was quadratic for rainbow trout, with a peak value at 24 to 36kg m⁻³.