Effect of microbial films on settlement of bryozoan larvae (Bugula simplex,B. stolonifera and B. turrita)

The influence of microbial films on larval settlement was studied for sympatric species of cellularioid cheilostome bryozoans (Bugula simplex, B. stolonifera, and B. turrita). Settlement experiments were conducted in the laboratory in 1981 with and without a choice of filmed or unfilmed substrates. When given a choice of substrates, the larvae of all three bryozoan species significantly preferred the filmed substrate. When not given a choice of substrates, both B. simplex and B. turrita larvae tended not to settle on the unfilmed substrate, while the filmed and unfilmed substrates were equally attractive to B. stolonifera. Microbial films enhanced the settlement of all three species.     

Influence du mode de conservation sur la composition chimique élémentaire de Pontella mediterranea (Copepoda: pontellidae)

A comparative study of the elementary chemical composition (C, H, N) of Pontella mediterranea (males, females and copepodites) was carried out in two series: in Series 1, live copepods were briefly rinsed with distilled water, dried for one night (12 h) at 60°C, and then stored in a desiccator; in Series 2, the copepods were preserved in a 5% formalin-freshwater solution neutralized with borax; they were then rinsed with distilled water, and subsequently dried and stored in a desiccator as in Series 1. The results reveal that the dry weight of preserved copepods (Series 2) is 20 to 25% less than that of fresh, dried individuals. Carbon loss was estimated as about 10% in males and 17% in females, hydrogen loss as 14% in males and 23% in females, and nitrogen loss as 20% in males and 21% in females. The organic compounds lost seem to be mainly proteinaceous. Nevertheless, relative carbon content expressed as percent dry weight, and C:N and C:H ratios were all significantly higher in the preserved specimens. These results are compared with literature data on two other crustancean species.     

Settlement of the tube worm Hydroides dianthus (Polychaeta: Serpulidae): cues for gregarious settlement

The larvae of many benthic marine invertebrates settle to form conspecific aggregations and are thought to rely on chemical cues associated with adults as indicators of habitat suitability, although the identification of inductive compounds has proven difficult. Still-water laboratory assays carried out during the summers of 1992 and 1993 with larvae of the serpulid polychaete, Hydroides dianthus (Verrill, 1873), demonstrate that unidentified water-borne compound(s) were responsible for gregarious settlement of competent larvae. Unlike inductive compounds associated with other tube-dwelling polychaetes, the settlement cue was soluble in water and was not associated with the tube, but rather with the body of live adults. In assay chambers divided by a 52-m mesh barrier, a greater percentage of larvae settled on biofilmed substrata when adult worms were present on the other side of the barrier than when adults were absent. Settlement in response to conspecific adults, live worms removed from their tubes, and amputated tentacular crowns of live worms was significantly greater than settlement in response to dead worms, empty tubes, or biofilmed slides. The settlement inducer appears to emanate from the openings of occupied tubes; settlement was greatest along the anterior two-fifths of the tube of living conspecific adults. A single adult was equally capable of eliciting a gregarious response as were five or 25 conspecifics, and newly settled juveniles began to elicit gregarious settlement after approximately 96 h. Extraction of aggregations of adult worms with organic solvents removed the inductive capacity of the tissue, and activity was found in both nonpolar and polar fractions of an extraction series.     

Influence of sponges on invertebrate recruitment: A field test of allelopathy

We manipulated live sponges in Belize, Central America, Big Pine Key, Florida (USA), and Indian River lagoon, Florida (USA) in summer/autumn, 1988. At each location, live sponges of three species were placed within 0.5 cm of ceramic tiles. Tiles with synthetic sponges positioned in the same manner and tiles with no sponges served as controls. Of 26 recruiting species analyzed, only one (Sponge sp. 6 — Indian River) was inhibited by living sponges. Four species (Perophora regina — Belize;Aiptasia pallida — Big Pine Key; andCrassostrea virginica andAscidia nigra — Indian River) recruited in greater numbers in the presence of sponges, suggesting that some larvae may be attracted rather than repelled by sponge allelochemicals. Allelopathic effects were less important than small-scale flow effects and patchy larval supply in determining recruitment patterns on surfaces adjacent to sponges.     

Choice and utilization of oviposition sites by female Calopteryx maculata (Odonata: Calopterygidae)

Summary Females of the damselfly Calopteryx maculata (de Beauvois) initially choose the larger of a pair of adjacent oviposition sites, about 70% of the time (Table 1), or whichever of two equal sized sites had other ovipositing females on it (about 88% of the time-Table 2). These criteria for initial choice between a pair of sites also interact. Incoming females generally (57 to 74% of the time-Table 3) joined others on the small site rather than ovipositing alone at the adjacent, bigger site. When pairs of large and small sites were replicated across eight locations, there were nonsignificant trends towards greater utilization (eggs laid) of the larger of a pair of oviposition sites within locations (Table 4). The lack of agreement between initial choice and utilization shows that other factors besides size are important in the choice and use of oviposition sites. These include disturbance by males, the presence of other females and choice criteria that can only be assessed during oviposition. When all sites at the eight locations were equal in size, there was considerable day to day and location to location variation in eggs laid (Fig. 1). Viewed over periods of several days, some sites are obviously less attractive than others in terms of cumulative numbers of eggs laid at them. When the amount of vegetation was varied among locations, those with the bigger oviposition sites were used more often, somtimes significantly so, but there were also significant reversals (small sites used more often) (Table 5, Fig. 1). Thus, there is no simple effect of size on the utilization of oviposition sites by Calopteryx maculata females, despite a clear tendency for females to make initial choices based on this criterion. The considerable among and within location variation in number of eggs laid may reflect additional choice criteria or the interaction of size, the presence of other females, disturbance, and location.     

The interplay between foraging mode, habitat structure, and predator presence in antlions

Antlion larvae are sand-dwelling insect predators, which ambush small arthropod prey while buried in the sand. In some species, the larvae construct conical pits and are considered as sit-and-wait predators which seldom relocate while in other species, they ambush prey without a pit but change their ambush site much more frequently (i.e., sit-and-pursue predators). The ability of antlion larvae to evade some of their predators which hunt them on the sand surface is strongly constrained by the degree of sand stabilization or by sand depth. We studied the effect of predator presence, predator type (active predatory beetle vs. sit-and-pursue wolf spider), and sand depth (shallow vs. deep sand) on the behavioral response of the pit building Myrmeleon hyalinus larvae and the sit-and-pursue Lopezus fedtschenkoi larvae. Predator presence had a negative effect on both antlion species activity. The sit-and-wait M. hyalinus larvae showed reduced pit-building activity, whereas the sit-and-pursue L. fedtschenkoi larvae decreased relocation activity. The proportion of relocating M. hyalinus was negatively affected by sand depth, whereas L. fedtschenkoi was negatively affected also by the predator type. Specifically, the proportion of individual L. fedtschenkoi that relocated in deeper sand was lower when facing the active predator rather than the sit-and-pursue predator. The proportion of M. hyalinus which constructed pits decreased in the presence of a predator, but this pattern was stronger when exposed to the active predator. We suggest that these differences between the two antlion species are strongly linked to their distinct foraging modes and to the foraging mode of their predators. Reut Loria and Inon Scharf contributed equally to the paper.     

Evidence for a water-borne cue inducing metamorphosis in the dorid nudibranch mollusc Adalaria proxima (Gastropoda: Nudibranchia)

The dorid nudibranch Adalaria proxima (Alder & Hancock) is a specialist predator of the cheilostome bryozoan Electra pilosa (L.). Natural induction of metamorphosis of the pelagic lecithotrophic larva of A. proxima was assessed in response to solutions from sonicated prey tissue and (live) E. pilosa-conditioned seawater (Electra-CSW). We exploited the tendency of larvae to become entrapped (rafted) at the air-water interface in cultures to examine whether larvae require direct contact with the live prey for metamorphosis to proceed. Larvae metamorphosed when rafted above colonies of live E. pilosa, above plankton mesh bags isolating live E. pilosa, and in choline chloride controls; there was no metamorphosis of larvae that were rafted in filtered seawater controls. Entrapped veliger shells remained rafted throughout the experimental period in all cases. No metamorphosis occurred in treatments containing either the supernatants or pelleted particulates obtained from sonicated colonies of E. pilosa. Both one-colony and three-colony Electra-CSW induced metamorphosis of larvae. These data are at variance with previous results in showing that direct contact with the live prey is not necessary for metamorphosis to proceed. Furthermore, the fact that competent larvae metamorphosed in response to Electra-CSW in the absence of any other cue strongly suggests that the inductive cue is water-borne.     

Substrate choice and settlement preferences of planula larvae of five Scyphozoa (Cnidaria) from German Bight,North Sea

The settlement behaviour of planula larvae and their development to young polyps was investigated in laboratory experiments in five scyphozoan species [Aurelia aurita (L.), Cyanea capillata (L.), Cyanea lamarckii Péron and Leseur, Chrysaora hysoscella (L.), and Rhizostoma octopus (L.)]. The undersides of settling plates were strongly preferred for settlement. Shells, the only natural substrate type offered, were less attractive than artificial substrates (concrete, machined wood, polyethylene, and glass). The advantages of colonization of substrate undersides for survival and reproduction of polyps are discussed. It is supposed that the increase of artificial substrates in our seas, due to marine litter pollution and submarine building activities, enlarge the areas of distribution of scyphozoan polyps, in coastal as well as in off-shore regions. Subsequent increases in ephyra production by polyps are probably one reason for the increase in mass occurrences of jellyfish recognized worldwide during the last few decades. It is suggested that the early developmental stages in the cnidarian life cycle, the planula larvae, and the polyps, play the key role in the development of jellyfish outbursts.     

Chemoecological studies of the exocrine glandular larval secretions of two chrysomelid species (Coleoptera):Phaedon cochleariae andChrysomela lapponica

Summary The exocrine glandular secretions of larvae of the subfamily Chrysomelinae are known to repel conspecific adults, other competitive phytophagous insects and natural enemies. InPhaedon cochleariae, the intraspecific activity of tlc fractions of the larval secretion was tested in order to examine the ecological significance of two fractions containing minor components and a fraction containing the major compound, the cyclopentanoid monoterpene (epi)chrysomelidial. InChrysomela lapponica, the defensive activity of the larval secretion against ants is known from specimens feeding upon willow or birch. The feeding preferences of larvae and adults ofC. lapponica from a Finnish and a Czech population were tested. The Finnish individuals significantly preferred feeding uponSalix borealis, whereas they hardly fed upon birch. The Czech specimens clearly preferred birch (Betula pubescens) to willow species. Application of salicin onto leaves of a willow species free of this phenolglycoside revealed that the Finnish individuals preferred feeding upon leaves with salicin. On the other hand, the Czech individuals avoided feeding upon leaves ofB. pubescens treated with salicin. The chemical composition of the glandular secretion of the Finnish larvae differed from the one of the Czech larvae. GC-MS-analyses of the secretions revealed that salicylaldehyde was the only major component of the secretion of Finnish larvae feeding upon the salicin-containing willowS. borealis. The glandular secretion of the Czech larvae feeding upon birch contained numerous esters of isobutyric acid and 2-methylbutyric acid. When Czech larvae had fed upon a salicin-containing willow (S. fragilis), the major compounds of their secretion were benzoic acid, salicylalcohol and benzoic acid esters; salicylaldehyde was only detected in traces. Thus,C. lapponica individuals from the Finland population adapted so closely to a salicincontaining willow that they clearly prefer this plant for food and that they obviously derive their main larval defensive compound (salicylaldehyde) from their host-plant.     

Polygyny,relatedness and nest founding in the polygynous myrmicine ant Leptothorax acervorum (Hymenoptera; Formicidae)

Summary There is high within-nest relatedness for functional queens (with corpora lutea), nonfunctional queens (without corpora lutea), and workers in polygynous nests of Leptothorax acervorum. The high functional queen relatedness suggests that young mated queens are adopted back to their mother nest. Functional queen relatedness does not change with the number of queens present in the nest, suggesting that the number of generations of queens, on average two to three, is rather stable. Worker relatedness decreases with increasing number of functional queens per nest (Tables 5, 6). The number of queens contributing offspring to the nest (mothers), estimated from worker and functional queen relatedness, is lower than the number of functional queens, particularly in highly polygynous nests. Estimates of number of mothers in monogynous nests indicate that these nests previously were polygynous (Table 7). There is no correlation between nest relatedness and distance between nests, and budding-off, if present, thus appears to be a rare mode of nest founding (Table 8). There are no indications of inbreeding in the two populations studied since the frequency of heterozygotes is as high as expected from random mating (Table 4). Most likely, polygyny is the rule in L. acervorum and serves to secure the presence of queens in the nest.     

Population biology of Gnathophausia longispina (mysidacea: Lophogastrida) from a central north pacific seamount

Large numbers of the lophogastrid mysid Gnathophausia longispina were discovered in 1984 over Southeast Hancock Seamount on the northern Hawaiian Ridge. Additional specimens were collected from the same area during subsequent surveys in 1985, 1987, and 1988 to obtain new information on the life history of this species. Estimates of growth suggest that G. longispina reaches sexual maturity by its second year. Young are brooded by the female for nearly a year and released in late winter-early spring. It is doubtful if G. longispina live longer than 2 yr or produce more than one brood. Most G. longispina are found within 100m above the seamount summit. The modal progression of size classes over time indicates that this species is dependent on local processes rather than recruitment from other areas to maintain its populations at the seamount.     

Zoogeography of fish parasites of the pearlside (Maurolicus muelleri), with genetic evidence of Anisakis simplex (s.s.) from the Mid-Atlantic Ridge

A total of 200 Maurolicus muelleri from the Mid-Atlantic Ridge (MAR) and the Norwegian Deep (ND) were studied for parasitic helminths. Two different metazoans were isolated from the MAR and five species from the ND. The predominant parasite species in both areas were tetraphyllidean cestode larvae (Scolex pleuronectis) and the anisakid nematode Anisakis simplex sensu stricto (s.s.), the latter being identified using genetic analysis of the internal transcribed spacer (ITS-1, ITS-2) and 5.8S regions of the rDNA. The parasite fauna of M. muelleri from the MAR was less species rich in comparison to ND, due to the deep-sea and oceanic environment. The digeneans Brachyphallus crenatus and Lecithaster confusus as well as the raphidascarid fish nematode Hysterothylacium aduncum were only collected from the ND. This can be explained either by the deep origin of the sampled fish specimens or the lack of suitable intermediate or final hosts in the region. Based on the frequent occurrence of A. simplex (s.s.) around the MAR and the ND, a pelagic life cycle is suggested at both localities, involving baleen and toothed whales as final and pelagic and mesopelagic fish and invertebrates as intermediate or paratenic hosts.     

Vibratory communication in the soil: pupal signals deter larval intrusion in a group-living beetle <Emphasis Type="Italic">Trypoxylus dichotoma</Emphasis>

Pupae of several insect species are known to generate air-borne sounds and/or substrate-borne vibrations, but the functions of the sounds/vibrations are not well understood. Here, we present the first evidence of vibratory communication between pupae and larvae of a group-living Japanese rhinoceros beetle Trypoxylus dichotoma which inhabits humus soil. The last-instar larvae of this beetle construct their own pupal cells to ensure normal pupation and eclosion. These cells are fragile and subject to damage from burrowing larvae because pupae and larvae co-inhabit the same patches of humus. In laboratory experiments, we demonstrated that pupal cells harboring live pupae were less likely to be broken by larvae than those harboring dead pupae. It was also demonstrated that pupae produced vibrations in response to larvae approaching the pupal cells. High-speed video and vibration analyses showed that pupae emitted 3–7 pulses at 1.3-s intervals by beating their pronotum against the inner wall of the pupal cell. The pupal vibration was of low frequency with a maximum energy at ≈ 100 Hz. The drumming behavior was more frequently observed in the presence of an approaching larva than in its absence. When pupal vibrations were played back near to vacant artificial pupal cells, these cells were rarely disturbed by the larvae. These results provide evidence that pupae generate vibrations to deter conspecific larvae, thereby preventing damage to the cells. This larval response to pupal vibrations may have evolved through preexisting anti-predator and/or sib-killing-avoidance behavior.     

Influence of limited starvation periods on growth and elemental composition (C,N,H) of Carcinus maenas (Decapoda: Portunidae) larvae reared in the laboratory

Zoea-1 larvae of Carcinus maenas L. (Decapoda: Brachyura: Portunidae) were from Helgoland in March 1984 and reared in the laboratory at 18°C through ecdysis. Dry weight (DW) and elemental composition of carbon (C), nitrogen (N), and hydrogen (H) were analyzed in newly hatched zoea-1, after different initial starvation periods, and in newly moulted zoea-2. Continually starved zoea-1 lost biomass and energy steadily, and logarithmic functions show best fit of empirical and predicted data. Biomass and energy equivalents of newly moulted zoea-2 are significantly correlated with starvation periods in the zoea-1, showing lower values with longer initial starvation. After about 25 to 34% individual biomass and energy losses, larvae exceed the point-of-noreturn (PNR), and do not recover or moult to the zoea-2, even if re-fed. When starvation ceases before the PNR, larvae moult to the zoea-2, and develop with lower average growth rates (AGR) after prolonged periods of initial food deprivation. The later larvae were re-fed, the less absolute amounts of DW, C, H, and individual energy, but more DW-related energy equivalents and N accumulated during subsequent feeding towards ecdysis. It is suggested that lipid, rather than protein, is the main source of energy controlling the maintenance of larval moult cycles. After lipid reserves are depleted, zoea-1 larvae live on body protein, and lose the ability to absorb and restore sufficient lipid if re-fed later than the PNR.Contribution to research project An-145/1-1 granted by the Deutsche Forschungsgemeinschaft (DFG)     

Reproductive success of nesting male longear sunfish (Lepomis megalotis peltastes)

Summary Although a few male longear sunfish nest solitarily, most territorial males excavate their nest in dense aggregations. The importance of sexual selection in the evolution of this breeding system was evaluated by 1) examining the influence of certain male and nest characteristics on spawning success and 2) by comparing the success of social and solitary nesters. Among group nesters, females spawned preferentially with males nesting early within a spawning period and occupying central nests. Male size and nest diameter were negatively correlated with nesting day and hence spawning success. However, female discrimination using these traits directly could not be demonstrated. The reproductive success of social and solitary males was compared by counting the larvae in their nests several days after spawning. Successful males (with larvae) were more likely than males without larvae to be encountered later in the breeding season and to nest in small groups or solitarily. Spawning period (of which there were 5 or 6 in a season) was significantly correlated with larval abundance in the nest, while male size was not. Social and solitary males were equally successful, but aggregations may nonetheless result from sexual selection: males unlikely to attract females may nest around more attractive males to steal fertilizations from them. High spawning synchrony may prevent attractive males from renesting elsewhere once other nesters aggregate around them.     

Hybridization within the species complex of the scleractinan coral Montastraea annularis

The morphologically variable reef coral previously known as Montastraea annularis (Ellis and Solander, 1786) has recently been separated into three species based on differences in morphology, behavior, allele frequencies and some life-history traits of Panamanian specimens. To further investigate the proposed reclassification and its conformity to the biological species concept we conducted reciprocal intra- and inter- specific fertilization experiments with gametes from each of the three species on Florida reefs. With one exception, self-fertilization rates were very low or zero. Within-species crosses resulted in production of planulae, as did all inter-species (hybrid) crosses, but there was much variation in fertilization success within each type of cross. In an experiment with separated gametes, hybrid crosses between M. annularis and M. franksi produced more larvae than within-species crosses for each species. Hybridization crosses between M. faveolata and the other two species produced fewer larvae than did within- M. faveolata crosses in the experiment with separated gametes, but many larvae resulted when the hybridizations were performed by mixing entire gamete bundles. Additional observations showed that M. franksi had 20% larger eggs and fewer eggs per gamete bundle than did the other two species and that it consistently spawned 1 to 1.5 h before the others, a potential temporal barrier to hybridization. These results indicate that there is no inherent pre-zygotic barrier to cross-fertilization among the three morphological species, although post-zygotic survival and fertility remain to be determined. The adherence of the proposed reclassification to the biological species concept requires further examination. Received: 16 April 1997 / Accepted: 17 June 1997     

Lethal effects of Northwest Atlantic Ocean isolates of the dinoflagellate, Scrippsiella trochoidea, on Eastern oyster (Crassostrea virginica) and Northern quahog (Mercenaria mercenaria) larvae

The thecate dinoflagellate Scrippsiella trochoidea is a cosmopolitan, bloom-forming alga that has been generally considered non-toxic. Here, we report that environmentally relevant cell densities (10⁴ cells mL⁻¹) of Scrippsiella trochoidea strains isolated from the Northwest Atlantic Ocean caused 100% mortality in Eastern oyster (Crassostrea virginica) larvae during 3-day exposures while parallel control larvae exhibited 100% survival. S. trochoidea also exhibited lethal effects on Northern quahog (Mercenaria mercenaria) larvae (70% mortality during 3-day exposure) but were non-toxic to juvenile fish (Cyprinodon variegates). The cultures of S. trochoidea were more lethal to Northern quahog larvae than ten other species of harmful algae, including the highly toxic species Cochlodinium polykrikoides. Scrippsiella trochoidea cultures within later stages of growth were more toxic than exponential growth stages to bivalve larvae, and the toxicity was dose dependent. Furthermore, toxicity was maintained in the cultures that were sonicated, boiled, and frozen as well as in resuspended residues of the culture but was significantly lower in cell-free culture media. Collectively, these results suggest that S. trochoidea causes mortality in bivalve larvae through a physicochemical rather than strictly chemical mechanism, such as clogging of larval feeding apparatuses by materials produced by S. trochoidea (e.g., lipids, extracellular polysaccharides, and/or cell debris) which accumulate as cells in culture or blooms age. This is the first report of the lethal effects of Scrippsiella trochoidea on shellfish larvae.     

Role of surface chemical signals in egg cannibalism and intraguild predation in ladybirds (Coleoptera: Coccinellidae)

Summary. The eggs of some ladybirds are known to be toxic to intraguild ladybird predators. However, this defence is of little value if the eggs are killed before their toxicity becomes apparent. The results presented in this paper indicate that chemicals on the surface of the eggs of two species of ladybirds signal the relative risk of cannibalism and intraguild predation. In Adalia bipunctata and Coccinella septempunctata, 87% of the chemicals are alkanes. Each species of ladybird is less reluctant to eat their own eggs than those of the other species. This asymmetry is to be expected because there is a greater risk to ladybirds from intraguild predation than cannibalism. Similar alkanes to those on the surface of the eggs of A. bipunctata are present in tracks left by larvae and on the elytra of the adults of this species. Those in the larval tracks deter females from ovipositing in patches of prey already being attacked by their larvae and those on the elytra are used in mate recognition. That different context dependent messages could be signalled by similar chemicals is an example of semiochemical parsimony. Received 25 February 2000; accepted 22 May 2000     

Substratum preferences in planula larvae of two species of scleractinian corals, Goniastrea retiformis and Stylaraea punctata

To test whether coral planulae recruit randomly to different coral reef habitats or have specific substratum preferences, the settling behavior of planulae from two shallow water coral species from Pago Bay, Guam (13°25.02N, 144°47.30E) were examined in the laboratory in June and July of 1995. Goniastrea retiformis is generally restricted to the shallow reef front (<10 m depth) in areas dominated by crustose coralline algae (CCA), while Stylaraea punctata is abundant on inner reef flats were CCA coverage is low and sand and carbonate rubble covered by biofilms is common. When presented with four substrata (1) carbonate rock scrubbed free of biofilm and dried as a control, (2) the CCA Hydrolithon reinboldii, (3) the CCA Peyssonelia sp., and (4) naturally conditioned carbonate rubble covered by a biofilm, G. retiformis larvae showed a significant preference for H. reinboldii, and S. punctata larvae for the carbonate biofilm treatment. The preference shown by S. punctata larvae for biofilmed surfaces did not diminish with increasing larval age up to 11 days. These results suggest that the larvae of both species are capable of habitat selection, and that the preferred substrata among those tested bears a relationship to the habitats in which adult colonies were found. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Competition with flies promotes communal breeding in the burying beetle,Nicrophorus tomentosus

Communal breeding through nest-sharing may benefit cooperating individuals indirectly, in increased inclusive fitness, or directly, when environmental constraints reduce the fitness of solitary breeders. Burying beetles provide extensive parental care and can breed either in pairs or in larger groups of unrelated males and females. Parentage of communally-reared broods is usually shared but is skewed in favor of the individuals of each sex that provide longer care. Females provide care longer than males, and two females are more likely to remain together in the brood chamber than two males are. Flies and other burying beetles are the major competitors for carcasses and this study suggests that it is competition with flies that promotes communal breeding inNicrophorus tomentosus On medium-size carcasses (35–40 g) the presence or absence of oviposition by flies had a significant effect on the size of the brood reared, and on large carcasses (55–60 g) the number of beetles present, two or four, had a significant effect on brood size. On both medium and large carcasses, pairs rearing broods on flyblown carcasses had fewer young than pairs on clean carcasses or foursomes on flyblown carcasses. There was a strong trend for an interaction effect between number of beetles and competition with flies (Table 1). Duration of parental care was not affected by competition with flies except for that of the first male to depart, which provided care longer on flyblown carcasses (Table 2). Pairs and foursomes were equally able to defend the carcass and brood from conspecific intruders and from larger intrudingNicrophorus orbicollis (Table 3).