Dynamics of external brooding in the sea anemone Epictis Epiactis prolifera

The sexually produced young of the externally brooding actinian Epiactis prolifera Verrill, 1869 are attached to the parent's column just above the base. A transitory brood groove may be formed around the limbus when the parent contracts. In the population studied on the coast of Sonoma County, California, USA, from 27 to 49% of the adult anemones were brooding at any time, the proportion being inversely related to seawater temperature and directly related to size of the anemones. A brood usually consisted of young of various sizes. Overall, number of young being brooded was directly related to parent size and inversely related to size of the brooded juveniles. The growth rate of juvenile anemones was inferred to average about a millimeter in pedal disc diameter per month. Juveniles dislodged from the parent when smaller than 4 mm in basal diameter apparently cannot survive in the intertidal zone. Juveniles which successfully make the transition to adulthood have, therefore, been brooded for at least 3 months. Fewer than 50% of the smallest juveniles survived to a size at which they could live independently. The transition to adulthood was accompanied by a mortality of 80%. The survival rate, from egg to adult, was calculated to be about 1% per season.This work was done in partial fulfillment of the requirements for the Ph.D. degree in the Department of Zoology, University of California, Berkeley, and was supported by a U.S. National Science Foundation Traineeship.     

Effect of intraspecific density on life history traits and population growth rate of Neanthes arenaceodentata (Polychaeta: Nereidae) in the laboratory

The effects of intraspecific density on life history traits and population dynamics of the nereid polychaete Neanthes arenaceodentata Moore were assessed in a laboratory experiment. Survival, growth, and fecundity were measured for one generation of worms at densities of 40, 80 and 160 worms per 840 cm² (1x, 2x, and 4x treatments, respectively). Density did not affect size (prior to pairing), percentage of worms paired, time to pairing, or size of mature paired males. Density did have a significant negative effect on survival, size of mature paired females, time to spawning, percentage of females that reproduced, and number of eggs per reproducing female. As density increased, mean survival was 90.0, 80.8 and 74.0%; mean size of mature females was 52.2, 49.2 and 48.1 segments; mean time to spawning was 100.6, 102.4 and 109.4 d; and mean fecundity was 881, 622, and 598 eggs per female, for 1x, 2x and 4x treatments, respectively. Increased density reduced the potential population growth rate, ; for a given rate of larval survivorship, was lower in the 2x and 4x treatments than the 1x treatment. Analysis of sensitivity of to changes in survivorship indicated that population growth rate at the highest density was sensitive to both changes in larval survivorship and the probability of producing a successful brood, although at low densities, was sensitive only to changes in larval survivorship. We attribute these density effects to aggressive bahavioral interactions between the worms, primarily the adults. This experiment identifies key life history traits that could be measured in future experiments to test population level responses of N. arenaceodentata populations to pollutants, both in the laboratory and in the field.Contribution No. 820 of the U.S. Environmental Protection Agency, Environmental Research Laboratory and Contribution No. 189 of the Marine Sciences Institute, University of Connecticut     

Brooding and development of <Emphasis Type="Italic">Anasterias minuta</Emphasis> (Asteroidea: Forcipulata) in Patagonia,Argentina

Brooding, embryonic and larval development, and the influence of environmental and biological factors in tidepool habitats were studied in the sea star, Anasterias minuta, at various sites along ~220 km of the Patagonian coast. This species has a benthic, lecithotrophic development that includes eight distinct developmental stages. A larval organ, the connection cord, is developed from a small preoral lobe at early stages of development and becomes larger and thinner at advanced stages. Fecundity and average egg size increased with female body size. The regression of log egg number to log sea-star size and weight at different sites had a slope significantly less than 3.0, resulting in negative allometry and indicating that brood capacity was limited in large females. Development was generally synchronous among sites, but varied within each brood at advanced stages, with more developed brooded larvae located at the periphery of the brood mass. Brooding was synchronous among various populations at different years and spatial scales, and extended over a period of 8 months. The highest proportion of brooding females occurred during May and June (austral winter). Juveniles were released mainly during September. The likelihood of finding brooding sea stars decreased with increasing sea water temperature, tidal height, and wave exposure, and increased with increasing body size. Both body size of brooding females and brooding rate were higher in the infralittoral fringe than at midlittoral levels. A revision of the current model of brooding behavior and development among forcipulate sea stars is given.     

Bird Communities of Prairie Uplands and Wetlands in Relation to Farming Practices in Saskatchewan

Abstract: Modern farm practices can vary in their emphasis on tillage versus chemicals to control weeds, and researchers know little about which emphasis has greater ecological benefits. We compared avifaunas of uplands and wetlands in four treatments: conventional farms, conservation farms (contrasting those that minimized frequency of tillage [minimum tillage] with those that eliminated chemical inputs [organic]), and restored or natural (wild) sites in Saskatchewan, Canada. Of 37 different upland bird species encountered during surveys, one made greater use of farms, four made greater use of wild sites, and the remaining species showed no preference. When all upland species were combined, higher relative abundance occurred on wild than on farm sites, and on minimum tillage than on conventional farms. Wild upland sites also had more species than did conventional farms. Of 79 different species encountered during surveys of wetlands and their margins, most had similar encounter probabilities among treatments, although seven were more common on either organic farms or wild sites. Higher relative abundances were documented in wetland habitat of wild sites and organic farms than of minimum tillage or conventional farms. Wetlands of wild sites had more species than did minimum tillage or conventional farms. Overall, in terms of both avifaunal density and diversity, small treatment effects could be ascribed to differences between conventional and conservation farms, whereas larger effects were due to differences between farms and wild sites. Wetlands were heavily used by birds in all treatments, suggesting high conservation priority regardless of context.     

Prodissoconch morphology is environmentally modified in the brooding bivalve Lasaea subviridis

Embryonic shell sculpture of intertidal and laboratory brooded individuals of the direct developer Lasaea subviridis was studied using scanning electron microscopy. Intertidally brooded individuals develop a distinct prodissoconch I (PI) and prodissoconch II (PII) of unusual morphologies. The PI is relatively small (50–100 m in length), oval in outline, lacks a radial sculpture and is restricted to the initial pitted zone of the prejuvenile shell. A larger PII is formed (510–680 m in length) and it is dominated by pronouced commarginal striae but also includes a belt of radial ridges extending from the PI/PII boundary. Laboratory brooded individuals kept constantly submerged do not develop the pronounced commarginal striae characteristic of PII. This implies that PII formation in L. subviridis is not a direct result of the brooding habit, and can be modified by environmental factors, which possibly include low-tide exposure. There is no single feature of external prodissoconch morphology that unambiguously indicates a direct development mode combined with mantle cavity brood protection. General guidelines are presented to help recognize this developmental mode, based on prodissoconch morphology. In comparing prodissoconch morphologies of brooding bivalves, the habitats as well as the phylogenies of species should be considered, especially in comparisons of intertidal and subtidal species.     

Sea-Cage Aquaculture, Sea Lice, and Declines of Wild Fish

Abstract: A sea cage, sometimes referred to as a net pen, is an enclosure designed to prevent farm fish from escaping and to protect them from large predators, while allowing a free flow of water through the cage to carry away waste. Farm fish thus share water with wild fish, which enables transmission of parasites, such as sea lice, from wild to farm and farm to wild fishes. Sea lice epidemics, together with recently documented population‐level declines of wild salmon in areas of sea‐cage farming, are a reminder that sea‐cage aquaculture is fundamentally different from terrestrial animal culture. The difference is that sea cages protect farm fish from the usual pathogen‐control mechanisms of nature, such as predators, but not from the pathogens themselves. A sea cage thus becomes an unintended pathogen factory. Basic physical theory explains why sea‐cage aquaculture causes sea lice on sympatric wild fish to increase and why increased lice burdens cause wild fish to decline, with extirpation as a real possibility. Theory is important to this issue because slow declines of wild fish can be difficult to detect amid large fluctuations from other causes. The important theoretical concepts are equilibrium, host‐density effect, reservoir‐host effect, and critical stocking level of farmed fish (stocking level at which lice proliferate on farm fish even if wild fish are not present to infect them). I explored these concepts and their implications without mathematics through examples from salmon farming. I also considered whether the lice‐control techniques used by sea‐cage farmers (medication and shortened grow‐out times) are capable of protecting wild fish. Elementary probability showed that (where W is the abundance of wild fish, W* is the prefarm abundance, F is the abundance of farm fish, and is the ratio of lice per farm fish to lice per wild fish). Declines of wild fish can be reduced by short growing cycles for farm fish, medicating farm fish, and keeping farm stocking levels low. Declines can be avoided only by ensuring that wild fish do not share water with farmed fish, either by locating sea cages very far from wild fish or through the use of closed‐containment aquaculture systems. These principles are likely to govern any aquaculture system where cage‐protected farm hosts and sympatric wild hosts have a common parasite with a direct life cycle.     

Brood sorting by ants: distributing the workload over the work-surface

Summary Leptothorax unifasciatus ant colonies occupy flat crevices in rocks in which their brood is kept in a single cluster. In artificial nests made from two glass plates sandwiched together, designed to mimic the general proportions of their nest sites in the field, such colonies arrange their brood in a distinct pattern. These patterns may influence the priority with which different brood are tended, and may therefore influence both the division of labour and colony demography. Different brood stages are arranged in concentric rings in a single cluster centred around the eggs and micro-larvae. Successively larger larvae are arranged in progressive bands away from the centre of the brood cluster. However, the largest and oldest brood items, the prepupae and pupae, are placed in an intermediate position between the largest and most peripheral larvae and the larvae of medium size. Dirichlet tessellations are used to analyze these patterns and show that the tile areas, the area closer to each item than its neighbours, allocated to each type of item increase with distance from the centre of the brood cluster. There is a significant positive correlation between such tile areas and the estimated metabolic rates of each type of brood item. The ants may be creating a domain of care around each brood item proportional to that item's needs. If nurse workers tend to move to the brood item whose tile they happen to be within when they have care to donate, they may apportion such care according to the needs of each type of brood. When colonies emigrate to new nests they rapidly recreate these characteristic brood patterns.     

Pollutant effects on the mixed function oxygenase (MFO) and reproductive systems of the marine polychaeteNereis virens

The cytochrome P-450 dependent mixed-function oxygenase (MFO) system ofNereis virens, found in the endoplasmic reticulum of the intestinal epithelium, has similar components and cofactor requirements when compared to other invertebrates. Worms exposed to food containing benzo(a)pyrene or a mixture of polychlorinated biphenyls (Aroclor 1254) had significantly higher MFO activity and cytochrome P-450 content than unexposed ones.N. virens from a site in Porland, Maine, USA, which was heavily contaminated with oil, and nearby clean reference sites were studied. The weight, MFO activity and presence or absence of gametes were determined. The MFO activity in worms from the oil-contaminated site was approximately six times that of worms from the reference sites; worms from the reference areas were six times larger than oil-contaminated worms. Worms from the oil-contaminated area lacked or had undeveloped gametes in spring, summer and fall. In contrast, many of the worms collected from the reference sites at the same times had fully developed gametes.     

Extra-pair paternity and intraspecific brood parasitism in wild zebra finches Taeniopygia guttata,revealed by DNA fingerprinting

Summary The frequency of extra-pair parentage in a wild population of zebra finches Taeniopygia guttata was examined by DNA fingerprinting. A total of 25 families, comprising 16 pairs of parents and 92 offspring (in broods of 1 to 6) were examined. Ten cases of extra-pair parentage, presumed to constitute intraspecific brood parasitism, were detected (10.9% of offspring or 36% of broods), including one possible instance of quasi-parasitism (parasitism by a female fertilized by the male nest owner). The average number of parasitic eggs per clutch detected by fingerprinting was 1.10±0.32 SD, very similar to the one egg difference in average clutch size between parasitised (6.0±0.82) and unparasitised nests (5.0±0.95). Two cases of extra-pair paternity (EPP) were detected among 82 offspring whose maternity was confirmed: 2.4% of offspring, or 8% of broods. In both cases EPP accounted for only a single offspring within a brood. Behavioural observations show that EPP occurs through extra-pair copulation rather than rapid mate switching. The results are discussed in the light of what is known about the fertile period and sperm precedence patterns in this species. Offprint requests to: T.R. Birkhead     

Green abalone, <Emphasis Type="Italic">Haliotis fulgens</Emphasis> infected with the agent of withering syndrome do not express disease signs under a temperature regime permissive for red abalone, <Emphasis Type="Italic">Haliotis rufescens</Emphasis>

All California abalone species have been shown to be susceptible to infection with the bacterial agent of abalone withering syndrome (WS), although expression of signs of the disease may vary between species and with environmental conditions. We examined thermal modulation of WS expression in green abalone Haliotis fulgens at temperatures mimicking El Niño (18.0°C) and La Niña (14.2°C) events in southern California. In contrast to results obtained from previous experiments with red abalone, H. rufescens, the higher temperature did not result in higher infection intensities of the causative agent of the disease nor increase in clinical signs of disease. These results demonstrate clear differences in thermal regulation of disease expression between abalone species, and provide further data suggesting that green abalone should be a target species of recovery efforts in southern California, where WS is endemic.     

Spirorbinae (Serpulidae: Polychaeta) from South Africa,including three new species

Five Antarctic or south-temperate species of Spirorbiaae are centred in south-west Africa, whilst 6 tropical species extend round the south-east and another 4 are recorded from Mozambique. Of the three new species, Pileolaria (Pileolaria) annectans sp. n. has a rudimentary opercular plate surmounting a sort-walled sac of embryos, which somewhat resembles a thoracic brood pouch. Pileolaria (Pileolaria) dayi sp. n. has a helmet-like brood chamber, but this opens distally, like the brood cups found in the subgenus Duplicaria. Janua Pil-natalensis sp. n. has the dorsal collar folds usually separate, but partially fused in some juveniles.     

Sex-specific energy requirements in nestlings of an extremely sexually size dimorphic bird, the European sparrowhawk (Accipiter nisus)

Allocation of parental investment is predicted to be equal at the population level between both sexes of offspring, and should lead to sex ratio biases in species that exhibit a sex-difference in parental care. Sex-differences in parental care are rarely quantified. We measured daily energy expenditure in free-living nestlings of the extremely sexually size dimorphic European sparrowhawk (Accipiter nisus), using the doubly labelled water method. These data were combined with measured growth characteristics to estimate daily and total metabolised energy intake of male and female young during the nestling stage. Females reached an asymptotic body mass 1.6 times higher than males. This resulted in a total metabolised energy an estimated 1.4 times higher for the nestling stage. Furthermore, we observed a decline in daily metabolised energy with an increase in brood size, which was significantly stronger for females than for males. These results are discussed in the context of Fishers equal allocation theory. Empirical evidence of a sex ratio bias at the end of parental care, with an overall excess of males, is lacking in this species. Consequently, our data do not support the idea of equal allocation between the sexes. The observed sex difference in daily metabolised energy in response to brood size may give scope for sex ratio bias at the level of the individual brood.     

Predicting Human-Carnivore Conflict: a Spatial Model Derived from 25 Years of Data on Wolf Predation on Livestock

Abstract:  Many carnivore populations escaped extinction during the twentieth century as a result of legal protections, habitat restoration, and changes in public attitudes. However, encounters between carnivores, livestock, and humans are increasing in some areas, raising concerns about the costs of carnivore conservation. We present a method to predict sites of human-carnivore conflicts regionally, using as an example the mixed forest-agriculture landscapes of Wisconsin and Minnesota (U.S.A.). We used a matched-pair analysis of 17 landscape variables in a geographic information system to discriminate affected areas from unaffected areas at two spatial scales (townships and farms). Wolves ( Canis lupus ) selectively preyed on livestock in townships with high proportions of pasture and high densities of deer ( Odocoileus virginianus ) combined with low proportions of crop lands, coniferous forest, herbaceous wetlands, and open water. These variables plus road density and farm size also appeared to predict risk for individual farms when we considered Minnesota alone. In Wisconsin only, farm size, crop lands, and road density were associated with the risk of wolf attack on livestock. At the level of townships, we generated two state-wide maps to predict the extent and location of future predation on livestock. Our approach can be applied wherever spatial data are available on sites of conflict between wildlife and humans.     

Particle-size selection by Pseudopolydora paucibranchiata (Polychaeta: Spionidae) in suspension feeding and in deposit feeding: influences of ontogeny and flow speed

Pseudopolydora paucibranchiata Okuda suspension feeds and deposit feeds at the sediment-water interface, where it is exposed to a variety of particles differing in physical characteristics and nutritional value. In flume experiments (conducted in August 1994 and May 1995) with two sizes of either suspended or deposited beads, I measured particle-size selection separately in each feeding mode. The same influences of palp width and of ambient flow speed were observed in each mode. At velocities 0.74 cm s^-1 there were no relationships between palp width and the proportion of gut contents composed of large beads. At velocities 1.8 cm s^-1 worms with narrower palps ingested relatively fewer large beads (and more small beads) than did worms with wider palps. Palp width and body length were linearly related, and results were similar when analyzed with body length as the independent variable. As flow speed increased, selectivity changed in a worm-size-specific manner: worms with a palp width -1. Assuming that in the field (1) particle size is the principle criterion for selection, and (2) the amount of digestible food component in deposited and suspended particles, respectively, is related to particle surface area and volume, I hypothesize that changes in selectivity as velocity rises can cause juveniles to experience a decreasing profitability of suspension feeding and a simultaneously increasing profitability of deposit feeding. Juveniles could maintain a diet of high food value despite flow variations by adjusting the proportion of time they spend suspension feeding relative to deposit feeding.     

Brood defense and age of young: a test of the vulnerability hypothesis

Summary In many altricial species including the great tit (Parus major) the intensity of brood defense against predators has been shown to increase with the age of the offspring. This effect has been ascribed amongst others to the young becoming more vulnerable as they age (vulnerability hypothesis). In a great tit population suffering heavy losses from brood depredation by the great spotted woodpecker (Dendrocopus major), we rendered first and second broods more vulnerable by artificially enlarging the entrance of the nest hole. Contrary to the vulnerability hypothesis, 16 experimental pairs defended their brood against a dummy great spotted woodpecker less vigorously than did 16 control pairs. Nest concealment behavior potentially compromising active defense was minimized by simultaneous playback of nestling distress calls, thus simulating the act of nest predation. This leaves the brood value hypothesis as an alternative functional explanation of the defense level — age effect. It predicts that parents should defend their brood in proportion to the reproductive value (or some more suitable cohortal equivalent measure) of their offspring. At present, this explanation pertains to one predator species. In first broods, but not in second broods, males defended them more vigorously than did their females. While this parallels previous experiments on brood defense against predators posing a much greater risk to the parents, two functional explanations previously put forward can hardly apply.     

Reproduction and sex reversal of the solitary vermetid gastropod<Emphasis Type="Italic"> Serpulorbis arenarius</Emphasis>

The reproductive biology of the Mediterranean vermetid gastropod Serpulorbis arenarius, a subtidal sessile solitary species that lives in shady rocky communities, was studied at two localities south-east of the Iberian Peninsula. Its population density is usually low, with specimens sometimes sparsely distributed. Its shell often appears to be covered by epibiotic organisms. The sex ratio is biased toward males (63%) in the populations of SE Spain, and the results point towards hermaphroditism with alternating sexuality. The results obtained suggest that all specimens are first males, change to females under some as yet unknown condition and, after the ovogenesis cycle has been completed, revert to the original sexual state (male). The possible adaptive forces that favour hermaphroditism in this species are discussed, and we hypothesise that the seizure of a spermatophore by males may stimulate sex change to female. Brooding females (51% of females studied) were found from March to December. In males, histological study of testes revealed that spermatogenesis seems to occur throughout the year. Females brooded between 5 and 28 (=15) egg capsules, which were attached by stalks to the inner surface of the shell. It is noteworthy that the gonads of two of the specimens brooding egg capsules were developing testes. The newly laid egg capsules were filled with an organic intracapsular matrix, in which 40–153 (=94) eggs were embedded. The mean diameter of the uncleaved ova was 352.5 µm. The organic intracapsular matrix and some nurse eggs were used as an extra-embryonic food source. Hatching occurred at an advanced veliger stage, immediately followed by metamorphosis. Therefore, the larval development of S. arenarius is of a lecithotrophic intracapsular type, with a short, free-swimming/crawling phase.Communicated by O. Kinne, Oldendorf/Luhe     

Population dynamics of three estuarine meiobenthic harpacticoids (Copepoda) in South Carolina

To elucidate the population dynamics of abundant meiofauna, a 15-month field study of 3 species of harpacticoid copepods was conducted in a South Carolina (USA) estuary. Per capita birth and death rates and average brood size (number of eggs per adult female) were lowest for Enhydrosoma propinguum (Brady), Microarthridion littorale (Poppe), and Stenhelia (Delavalia) bifidia (Coull) from November–February, indicating that food or density-independent factors such as temperature regulated winter densities. Reproduction for all species began in February, with the first appearance of copepodites approximately 1 month later. From March–October, M. littorale displayed high per capita birth and death rates (>0.6 individual^-1 day^-1), and high average brood size (8 to 9 eggs per adult female). As this species is epibenthic, and thus susceptible to grazing by detritivores, these data suggest that M. littorale was limited by predation. S. (Delavalia) bifidia displayed a sharp increase in density, birth rate and average brood size in July, but birth and death rates (0.3 individual^-1 day^-1) were always lower than those of M. littorale and average brood size gradually decreased from July–December. As this species burrows and is less susceptible to grazing predation, these data are consistent with the hypothesis that S. (Delavalia) bifidia became food-limited as population growth approached its carrying capacity. Although E. propinguum was the most abundant species, quantification of birth potential was not accomplished due to an underestimate of the number of ovigerous females. Therefore, little can be concluded about the mechanism of the summer density regulation of E. propinguum.Contribution No. 274 from the Belle W. Baruch Institute for Marine Biology and Coastal Research, University of South Carolina.     

Effects of pressure,temperature and oxygen on the oxygen consumption rate of the Midwater copepod Gaussia princeps

In this study we assessed the amount of lead accumulated in the body of a grazing mollusc by transfer from its algal food in laboratory experiments, and compared these results with the amounts found in naturally occurring molluscs and seaweed. Near La Jolla, California (USA), where the concentration of lead in seawater is probably less than 0.08 g l^-1, most of the naturally occurring Egregia laevigata contains less than 0.4 g Pb g^-1 wet weight. The total body masses, without shells, of juvenile Haliotis rufescens fed on this seaweed for 3 to 6 months showed similar concentrations. When, however, E. laevigata is placed for 1 to 6 days in seawater to which lead has been added (0.1 or 1.0 mg l^-1) both the seaweed and the abalone subsequently fed with it accumulate proportionally larger amounts of lead. After 6 months, young abalone fed on E. laevigata pretreated with 1.0 mg Pb l^-1 accumulated up to 21 g Pb g^-1 wet weight. This amount of lead had no apparent consequences on the growth or activity of the molluscs. Analyses of 6 different organs from adult abalone showed that the lead was selectively concentrated in the digestive gland. In the foot (muscle tissue), which is the part normally consumed by humans, only negligible amounts were found.     

Task specialization in a wild bee,Apis florea (Hymenoptera: Apidae), revealed by RFLP banding

Workers in a wild in situ colony of the dwarf honey bee, Apis florea, were observed undertaking the following behavior: liquid foraging, pollen foraging, guarding, stinging, fanning and wagging abdomen. Bees of each behavioral class were separately collected and frozen. Collections were made over a period of 10 days. Random samples of brood and workers were also collected. DNA was extracted from each bee and fingerprinted using a probe of unknown sequence obtained from an A. mellifera genomic library. Patterns of fingerprints (Fig. 1) were dissimilar among behavioral classes (Tables 1 and 2), strongly suggesting a genetic component to division of labor in this species. This result supports similar findings in A. mellifera in a species that is not troubled by many of the experimental difficulties inherent in A. mellifera. Correspondence to: B.P. Oldroyd     

The biology of a subtropical population of Halictus ligatus Say (Hymenoptera: Halictidae)

Summary A large population of Halictus ligatus was studied in the subtropical climate of Knights Key, Monroe County, Florida. The dissection of 858 female bees caught on flowers and 420 bees from completely excavated nests gives the following picture of phenology, colony development and social organisation. In the Florida keys, H. ligatus is continuously brooded and multivoltine. However, towards the coldest time of year young gynes may rest in their natal nests rather than found a new colony. This may result in a partial synchronisation of nest initiation when warm weather returns after a particularly cold spell. Most nests are started by a single foundress that usually survives until near the end of the production of reproductives. The first brood is very variable in size and males average 11% of the bees produced at this stage. This figure increases to 56% when the first brood workers begin provisioning. Queens are produced some time after the rise in male production and colony longevity is extended by the presence of some worker brood during this phase. Queens average 16% larger than their workers but appear to exert little inhibition of worker reproductivity: 57% of worker bees mate and 68% show ovarian development. This population is unique amongst social halictines in being continuously brooded, multivoltine and in having such weak physiological caste differentiation. It seems to represent an intermediate stage between the primitively eusocial colonies of H. ligatus found in temperate regions and the communal-like ones of the tropics.