Antennal responses to floral scents in the butterflies <Emphasis Type="Italic">Inachis io</Emphasis>, <Emphasis Type="Italic">Aglais urticae</Emphasis> (Nymphalidae), and <Emphasis Type="Italic">Gonepteryx rhamni</Emphasis> (Pieridae)

Summary. To better understand the biological role of floral scents for butterflies, electrophysiological responses to floral scents were investigated using combined gas chromatography and electroantennographic detection (GC-EAD). The antennal responses of three butterfly species, Aglais urticae L. (Nymphalidae), Inachis io L. (Nymphalidae), and Gonepteryx rhamni L. (Pieridae) to floral scent compounds from both natural and synthetic mixtures were examined. Floral scents were collected from the butterfly nectar plants Cirsium arvense (L.) (Asteraceae), and Buddleja davidii Franchet cv. (Loganicaeae) with dynamic head-space methods on Tenax-GR and eluted with pentane. These eluates, composed of natural floral scent blends, represent an array of compounds in their natural state. In the GC-EAD analyses eleven compounds were identified from C. arvense with the benzenoid compound phenylacetaldehyde in highest abundance. Seventeen compounds were identified from B. davidii with the irregular terpene oxoisophorone in highest abundance. Thirty-nine synthetic floral scent compounds were mixed in pentane, in equal amounts; about 35 ng were allowed to reach the antennae. The butterflies showed antennal responses to most of the floral scent compounds from both natural and synthetic blends except to the highly volatile monoterpene alkenes. Certain benzenoid compounds such as phenylacetaldehyde, monoterpenes such as linalool, and irregular terpenes such as oxoisophorone, were emitted in relatively large amounts from C. arvense and B. davidii, and elicited the strongest antennal responses. These compounds also elicited strong antennal responses when present in the synthetic scent blends. Thus, the butterflies seem to have many and /or sensitive antennal receptors for these compounds, which points to their biological importance. Moreover, these compounds are exclusively of floral scent origin. For B. davidii, which depends highly on butterflies for pollination, the exclusive floral scent compounds emitted in high abundance could be the result of an adaptive pressure to attract butterflies. Received 2 Septemter 2001; accepted 9 September 2002.     

Foraging responses in the butterflies <Emphasis Type="Italic">Inachis io</Emphasis>, <Emphasis Type="Italic">Aglais urticae</Emphasis> (Nymphalidae), and <Emphasis Type="Italic">Gonepteryx rhamni</Emphasis> (Pieridae) to floral scents

Summary. For butterflies to be efficient foragers, they need to be able to recognize rewarding flowers. Flower signals such as colours and scents assist this recognition process. For plant species to attract and keep butterflies as pollinators, species-specific floral signals are crucial. The aim of this study is to investigate foraging responses to floral scents in three temperate butterfly species, Inachis io L. (Nymphalidae), Aglais urticae L. (Nymphalidae), and Gonepteryx rhamni L. (Pieridae), in behavioural choice bioassays. The butterflies were allowed to choose bet-ween flower models varying in scent and colour (mauve or green). Flowers or vegetative parts from the plants Centaurea scabiosa L. (Asteraceae), Cirsium arvense (L.) (Asteraceae), Knautia arvensis (L.) (Dipsacaceae), Buddleja davidii Franchet (Loganicaeae), Origanum vulgareL. (Lamiaceae), Achillea millefolium L. (Asteraceae), and Philadelphus coronarius L. (Hydrangiaceae) were used as scent sources. All visits to the models — those that included probing and those that did not — were counted, as was the duration of these behaviours. Both flower-naive and flower-experienced (conditioned to sugar-water rewards, the colour mauve, and specific floral scents) butterflies were tested for their preference for floral versus vegetative scents, and to floral scent versus colour. The butterflies were also tested for their ability to switch floral scent preferences in response to rewards. Flower-naive butterflies demonstrated a preference for the floral scent of the butterfly-favourable plants C. arvense and K. arvensis over the floral scent of the non-favourable plants Achillea millefolium (Asteraceae), and Philadelphus coronarius cv. (Hydrangiaceae). Most of the butterflies that were conditioned to floral scents of either C. arvense, K. arvensis, or B. davidii readily switched theirfloral scent preferences to the one most recently associated with reward, thus demonstrating that floral scent constancy is a result from learning. These findings suggest that these butterflies use floral scent as an important cue signal to initially identify and subsequently recognize and distinguish among rewarding plants. Received 2 September 2001; accepted 9 September 2002.     

Pyrrolizidine alkaloid content in <Emphasis Type="Italic">Senecio</Emphasis>: ontogeny and developmental constraints

Summary. The ontogeny of pyrrolizidine alkaloid (PA) synthesis and constraints on defence level during the seedling stage were examined in the annual Senecio vulgaris and the monocarpic perennial Senecio jacobaea. In both species, PAs were actively synthesized from the onset of seedling growth so that juvenile stages did not go through an undefended stage. Roots are known to be the exclusive sites where PAs are produced. Root biomass was the single most important biomass parameter explaining variation in total PAs per seedling. All correlation coefficients between—relative growth rate and PA concentration were negative, but none was significant. However, a significant negative—correlation was found between shoot to root ratio and PA concentration in S. jacobaeaseedlings, suggesting a dilution effect of the PAs. Earlier studies have shown that the shoot to root ratio is positively correlated with relative growth—rate of established S. jacobaea plants. It is therefore suggested that young S. jacobaea plants with a high shoot to root ratio and hence a high growth capacity necessarily have lower PA defence levels than plants with a low shoot to root ratio. Received 10 July 2002; accepted 16 November 2002.     

Uptake and metabolism of [<Superscript>14</Superscript>C]rinderine and [<Superscript>14</Superscript>C]retronecine in leaf-beetles of the genus <Emphasis Type="Italic"> Platyphora</Emphasis> and alkaloid accumulation in the exocrine defensive secretions

Summary. Sequestration and processing of pyrrolizidine alkaloids (PAs) by leaf beetles of the genus Platyphora were investigated. Tracer experiments with labeled alkaloids were performed with P. eucosma feeding on Koanophyllon panamense (Asteraceae, tribe Eupatorieae). P. eucosma catalyzes the same reactions previously demonstrated for P. boucardi specialized to Prestonia portobellensis (Apocynaceae): (i) epimerization of rinderine to intermedine; (ii) esterification of retronecine yielding insect-specific PAs; (iii) efficient transport of the PAs as free bases into the defensive secretions. P. bella feeding on Tournefortia cuspidata (Boraginaceae) shows the same sequestration behavior and ability to synthesize the specific retronecine esters. P. ligata, a species phylogenetically closely related to the PA adapted species and clustering in the same clade, but feeding on a host plant devoid of PAs, feeds easily on PA treated host-plant leaves, but does not sequester or metabolize PAs. P. kollari a species clustering outside the PA clade refused to feed on its food-plant leaves painted with PAs. The results are discussed in relation to host-plant selection of the PA adapted species and the role of PAs in chemical defense. Received 20 September 2002; accepted 18 November 2002.     

Pheromone-based aggregation behaviour of the firebrat, <Emphasis Type="Italic">Thermobia domestica</Emphasis> (Packard) (Thysanura: Lepismatidae)

Summary. We tested the hypothesis that aggregation behaviour of the firebrat, Thermobia domestica (Packard) (Thysanura: Lepismatidae), an inhabitant of enclosed microhabitats, is mediated, at least in part, by a pheromone. Individual insects were released into the central chamber of a 3-chambered olfactometer and test stimuli were placed in lateral chambers. Paper discs previously exposed for 3 days to 10 female, male, or juvenile T. domestica were all preferred by female, male, or juvenile T. domestica over unexposed paper discs, indicating the presence of an aggregation/arrestment pheromone. In additional experiments, frass and scales from female T. domestica, tested singly and in combination, proved not to be the source of the pheromone. Physical contact was required for pheromone recognition, indicating that the pheromone arrests rather than attracts conspecifics. Arrestment by the long-tailed silverfish, Ctenolepisma longicaudata Escherich (Thysanura: Lepismatidae), but not by the common silverfish, Lepisma saccharina L. (Thysanura: Lepismatidae), to T. domestica exposed paper discs suggests closer phylogenetic relatedness between C. longicaudata and T. domestica, than between C. longicaudata and L. saccharina. Whether C. longicaudata or L. saccharina produce an aggregation signal, and whether T. domestica respond to this signal is unknown. Received 10 June 2002; accepted 30 September 2002.     

Comparative sensitivity to and consumption of methyl eugenol in three <Emphasis Type="Italic">Bactrocera dorsalis</Emphasis> (Diptera: Tephritidae) complex sibling species

Summary. Sensitivities to methyl eugenol of three sibling species in the Bactrocera dorsalis complex were compared. The degree of species sensitivity to methyl eugenol, i.e. B. dorsalis > B. papayae > B. carambolae (in decreasing order), was concomitant with the species age-related response to methyl eugenol as previously reported. The ability to consume methyl eugenol by the three sibling species showed similar trend - the average ME consumption per male was 0.70 ml for B. dorsalis, 0.58 ml B. papayae and 0.18 μl B. carambolae. Results obtained were discussed in relation to area-wide control of fruit fly. Received 21 April 2002; accepted 9 July 2002     

Cardiac output during exposure to <Emphasis Type="Italic">Chattonella</Emphasis><Emphasis Type="Italic">marina</Emphasis> and environmental hypoxia in yellowtail (<Emphasis Type="Italic">Seriola</Emphasis><Emphasis Type="Italic">quinqueradiata</Emphasis>)

The experiments were undertaken to measure, for the first time, cardiac output in yellowtail (Seriola quinqueradiata) during exposure to a harmful red tide flagellate (Chattonella marina). The responses were compared with those during exposure to environmental hypoxia to evaluate the significance of the drop of arterial oxygen partial pressure (PaO₂) in the fish-kill mechanisms by C. marina. PaO₂ immediately decreased, whereas heart rate (HR) was maintained until shortly before death during exposure to C. marina. Suffocation developing during the exposure resulted from a decrease in blood oxygen content, but not from lowered blood flow to the tissue. Although exposure to both C. marina and hypoxia immediately decreased PaO₂, arterial oxygen content (CaO₂) and pH (pHa) were significantly lower, whereas HR and cardiac output (&Qdot;) remained significantly higher, for the C. marina-exposed fish than for hypoxia-exposed fish. Although the drop in PaO₂ appears to play a pivotal role in the mechanisms of fish death by C. marina, other physiological response(s) should also be considered.     

Mate preferences in <Emphasis Type="Italic">Drosophila</Emphasis> infected with <Emphasis Type="Italic">Wolbachia</Emphasis>?

The maternally inherited bacterium Wolbachia pipientis generates strong reproductive incompatibilities between uninfected females and infected males (cytoplasmic incompatibility), significantly reducing both female and male reproductive success. Such fitness costs are thought to place selective pressure on hosts to evolve pre-copulatory preferences for mating with compatible mates, thereby enabling them to avoid the reproductive incompatibilities associated with Wolbachia. Therefore, uninfected females are predicted to prefer mating with uninfected males, whereas infected males are predicted to prefer mating with infected females. Despite these predictions, previous investigations of pre-copulatory mate preferences in Wolbachia-manipulated Drosophila have not found evidence of female preference for uninfected or compatible males. However, none of these studies utilised a design where focal individuals are provided with a simple choice in a relatively non-competitive situation. We examined both female and male pre-copulatory mate preference based on mate infection status in Drosophila simulans and D. melanogaster using simple choice assays involving between 30–50 replicates per treatment. Although we found no evidence of female pre-copulatory mate preferences in either species, male D. simulans exhibited some preference for mating with females of the same infection status. However, this preference was not evident when we repeated the experiment to confirm this finding. Consequently, we conclude that neither male nor female D. melanogaster and D. simulans exhibit significant Wolbachia-associated pre-copulatory mate preferences.     

Effects of substrate type on growth and mortality of blue mussels (<Emphasis Type="Italic">Mytilus</Emphasis><Emphasis Type="Italic">edulis</Emphasis>) exposed to the predator <Emphasis Type="Italic">Carcinus</Emphasis><Emphasis Type="Italic">maenas</Emphasis>

Structure and complexity of the substrate are important habitat characteristics for benthic epifauna. The specific growth and mortality rates and inducible defence characters on medium-sized blue mussels (Mytilus edulis L.) exposed to shore crabs (Carcinus maenas L.) were examined on three different substrate types in combined field and laboratory experiments. The experiments showed that complexity of the substrate increased blue mussel survival significantly, through a decrease in predation pressure. However, increased intraspecific competition for food on the complex substrate resulted in significantly lower growth rates of the mussels. Inducible defence characters were also influenced by substrate type. Blue mussels were more affected by predators on the structurally simple substrate, where they developed thicker shells and a larger posterior adductor muscle.     

Individual specialization in the hunting wasp <Emphasis Type="Italic">Trypoxylon</Emphasis> (<Emphasis Type="Italic">Trypargilum</Emphasis>) <Emphasis Type="Italic">albonigrum</Emphasis> (Hymenoptera,Crabronidae)

Individual-level variation in resource use occurs in a broad array of vertebrate and invertebrate taxa and may have important ecological and evolutionary implications. In this study, we measured the degree of individual-level variation in prey preference of the hunting wasp Trypoxylon albonigrum, which inhabits the Atlantic Forest in southeastern Brazil. This wasp captures several orb-weaving spider genera to provision nests. Individuals consistently specialized on a narrow subset of the prey taxa consumed by the population, indicating the existence of significant individual-level variation in prey preferences. The population niche was broader in the wet season in terms of both prey size and taxa. In the case of prey size, the population niche expansion was achieved via increased individual niche breadths, whereas in the case of prey taxa, individual niches remained relatively constrained, and the population niche expanded via increased interindividual variation. The observed pattern suggests the possibility of functional trade-offs associated with the taxon of the consumed prey. The nature of the trade-offs remains unknown, but they are likely related to learning in searching and/or handling prey. We hypothesize that by specializing on specific prey taxa, individuals increase foraging efficiency, reducing foraging time and ultimately increasing reproductive success.     

Weaning in the guinea pig (<Emphasis Type="Italic">Cavia aperea</Emphasis> f. <Emphasis Type="Italic">porcellus</Emphasis>): Who decides and by what measure?

Offspring should be selected to influence maternal effort to maximize their own fitness, whereas mothers are selected to limit investment in present progeny. In mammals, this leads to a conflict over the amount of milk provided and the timing of weaning. The intensity and time course of such conflict has so far mostly been investigated experimentally in altricial rodents. However, it is expected that offspring options for conflict will depend on developmental state. We therefore investigated in the highly precocial domestic guinea pig (Cavia aperea f. porcellus) who decides over nursing performance and weaning and how pup state influences these decisions. Specifically, we tested whether a threshold mass of pups predicts weaning time. By exchanging older litters against neonates and vice versa, we produced a situation in which females differed in lactational stage from the cross-fostered pups. Our results indicate that females decide about the timing of weaning, as cross-fostered younger pups were weaned at a much younger age than controls and older pups benefited from continuing lactation of foster mothers. Growth rates did not differ in the treatment groups, and different weaning ages resulted in differing weaning mass refuting the hypothesis that weaning is based on a threshold mass of offspring. This constitutes clear evidence that in a precocial rodent, the guinea pig, decisions about maternal care are primarily determined by maternal state and little influenced by pup state despite the extreme precociality of offspring. We suggest that precocial pups show little resistance to early weaning when food is abundant, as they reach sufficient nutritional independence by the middle of lactation to enable independent survival.     

<Emphasis Type="Italic">Ceroplastes albolineatus</Emphasis>, the first scale insect shown to sequester pyrrolizidine alkaloids from its host-plant <Emphasis Type="Italic">Pittocaulon praecox</Emphasis>

Summary. The occurrence of pyrrolizidine alkaloids (PAs) in Pittocaulon (ex Senecio) praecox (Asteraceae) a species endemic to Mexico was established. The aboveground plant organs contain the 1,2-saturated monoester 7-angeloyl-dihydroxyheliotridane together with a small proportion of its 9-angeloyl isomer as major alkaloid. The monoesters are accompanied by the macrocyclic otonecine derivative senkirkine. Roots contain only related macrocyclic PAs with senecionine, senkirkine and platyphylline as major components; monoesters are absent. The broom-like succulent stems of P. praecox are infested by the scale insect Ceroplastes albolineatus conspicuously visible by its huge wax cover. All life-history stages, i.e. females, eggs, first instar nymphs (crawlers) and the wax cover were found associated with PAs. The measured PA concentrations clearly indicate sequestration. The highest PA concentrations (mg / g dry weight) reached are: mature females, 0.44; eggs, 0.58; crawlers, 0.37; wax cover, 0.08. The host plant as well as in the infesting scale insect contain the PAs exclusively as free bases. As a phloem-feeder C. albolineatus must acquire the PAs with the ingested phloem sap. This appears plausible since in Senecio species PA are transmitted and circulated through the phloem path. It is suggested that PAs may protect particularly the crawlers as the most endangered stage in the life-cycle of the scale insect.     

Environmental conditions influence egg color of reed warblers <Emphasis Type="Italic">Acrocephalus scirpaceus</Emphasis> and their parasite,the common cuckoo <Emphasis Type="Italic">Cuculus canorus</Emphasis>

The outer layer of the eggshell in birds is in many cases covered by pigments that are assumed to be genetically determined traits with a negligible environmental component. To test the hypothesis that spring environmental conditions (i.e., temperature and rainfall) may affect bird egg pigmentation, we measured by spectrophotometry and photography egg coloration and spottiness on reed warbler (Acrocephalus scirpaceus L.) clutches parasitized by the common cuckoo (Cuculus canorus L.) collected over a period of 24 years and preserved in the Zoological Museum, Copenhagen, Denmark. In addition, we investigated whether spring environmental conditions may influence the coevolutionary relationship between the cuckoo and its host via changes in cuckoo–host egg matching. Generalized mixed models revealed that reed warbler eggs were more brilliant in those springs with a higher rainfall and tended to be bluer and greener in springs with a lower relative temperature. On the other hand, cuckoo eggs were bluer and greener in springs with a higher rainfall. Cuckoo–host egg matching in blue-greenness and spottiness was better in springs with a higher rainfall. These results provide support for the existence of an environmental component on bird egg coloration and suggest that environmental factors may potentially affect the outcome of important features of the arms race between cuckoos and reed warblers.     

Altered c-Fos expression demonstrates neuronal stress in mummichog, <Emphasis Type="Italic">Fundulus heteroclitus</Emphasis>, exposed to <Emphasis Type="Italic">Pfiesteria shumwayae</Emphasis> and <Emphasis Type="Italic">Chaetoceros concavicornis</Emphasis>

To better understand sublethal effects of harmful algal blooms (HABs) on fish, mummichog, Fundulus heteroclitus (L.), were exposed in the laboratory to varying, environmentally relevant densities of Pfiesteria shumwayae (Glasgow et Burkholder, CCMP 2089, dinoflagellate) and Chaetoceros concavicornis (Mangin, CCMP 169, diatom). Two experiments were conducted during the spring of 2003 and 2004 to quantitatively examine the effects of acute (2 h) P. shumwayae and C. concavicornis algal exposure on mummichog brain activity using c-Fos expression as a marker of altered neuronal activity. Brains from HAB-exposed fish were removed, sectioned, and stained using immunocytochemistry prior to quantifying neuronal c-Fos expression. Fish exposed to P. shumwayae and C. concavicornis showed increased c-Fos expression compared to unexposed control fish. A significant dose-response relationship was observed, with increased labeling in brains of fish exposed to higher cell densities for both HAB species tested (P ≤ 0.01). Increased labeling was found in the telencephalon, optic lobes, midbrain, and portions of the medulla. The greatest increases in expression were observed in the telencephalon of P. shumwayae-exposed fish, and in the telencephalon and optic lobes of C. concavicornis-exposed fish (P ≤ 0.01). These increases in c-Fos expression are consistent with other physical and chemical stress exposures observed in fish. Neuronal stress, evidenced by c-Fos expression, demonstrates a sublethal effect of exposure and changes in brain activity in fish exposed to HAB species.     

Inter-polyp genetic and physiological characterisation of <Emphasis Type="Italic">Symbiodinium</Emphasis> in an <Emphasis Type="Italic">Acropora valida</Emphasis> colony

Corals harbouring genetically mixed communities of endosymbiotic algae (Symbiodinium) often show distribution patterns in accordance with differences in light climate across an individual colony. However, the physiology of these genetically characterised communities is not well understood. Single stranded conformation polymorphism (SSCP) and real time quantitative polymerase chain reaction (qPCR) analyses were used to examine the genetic diversity of the Symbiodinium community in hospite across an individual colony of Acropora valida at the spatial scale of single polyps. The physiological characteristics of the polyps were examined prior to sampling with a combined O₂ microelectrode with a fibre-optic microprobe (combined sensor diameter 50–100 μm) enabling simultaneous measurements of O₂ concentration, gross photosynthesis rate and photosystem II (PSII) quantum yield at the coral surface as a function of increasing irradiances. Both sun- and shade-adapted polyps were found to harbour either Symbiodinium clade C types alone or clades A and C simultaneously. Polyps were grouped in two categories according to (1) their orientation towardps light, or (2) their symbiont community composition. Physiological differences were not detected between sun- and shade-adapted polyps, but O₂ concentration at 1,100 μmol photons m⁻² s⁻¹ was higher in polyps that harboured both clades A and C symbionts than in polyps that harboured clade C only. These results suggest that the acclimatisation of zooxanthellae of individual polyps of an A. valida colony to ambient light levels may not be the only determinant of the photosynthetic capacity of zooxanthellae. Here, we found that photosynthetic capacity is also likely to have a strong genetic basis and differs between genetically distinct Symbiodinium types.     

Seasonal variations in the growth rates of euphausiids (<Emphasis Type="Italic">Thysanoessa inermis</Emphasis>, <Emphasis Type="Italic">T. spinifera</Emphasis>, and <Emphasis Type="Italic">Euphausia pacifica</Emphasis>) from the northern Gulf of Alaska

The euphausiids Thysanoessa inermis (Kroyer 1846), Thysanoessa spinifera (Holmes 1900), and Euphausia pacifica (Hansen 1911) are key pelagic grazers and also important prey for many commercial fish species in the Gulf of Alaska (GOA). To understand the role of the euphausiids in material flows in this ecosystem their growth rates were examined using the instantaneous growth rate (IGR) technique on the northern GOA shelf from March through October in 2001–2004. The highest mean molting increments (over 5% of uropod length increase per molt) were observed during the phytoplankton bloom on the inner shelf in late spring for coastal T. inermis, and on the outer shelf in summer for T. spinifera and more oceanic E. pacifica, suggesting tight coupling with food availability. The molting rates were higher in summer and lower in spring, for all species and were strongly influenced by temperature. Mean inter-molt periods calculated from the molting rates, ranged from 11 days at 5°C to 6 days at 8°C, and were in agreement with those measured directly during long-term laboratory incubations. Growth rate estimates depended on euphausiid size, and were close to 0 in early spring, reaching maximum values in May (0.123 mm day⁻¹ or 0.023 day⁻¹ for T. inermis) and July (0.091 mm day⁻¹ or 0.031 day⁻¹ for T. spinifera). The growth rates for E. pacifica remained below 0.07 mm day⁻¹ (0.016 day⁻¹) throughout the season. The relationship between T. inermis weight specific growth rate (adjusted to 5°C) and ambient chlorophyll-a concentration fit a Michaelis–Menten curve (r ² = 0.48) with food saturated growth rate of 0.032 day⁻¹ with half saturation occurring at 1.65 mg chl-a m⁻³, but such relationships were not significant for T. spinifera or E. pacifica.     

Evidence of a spawning area of <Emphasis Type="Italic">Anguilla</Emphasis><Emphasis Type="Italic">marmorata</Emphasis> in the western North Pacific

Leptocephali of the tropical eel Anguilla marmorata have been consistently collected in the same area of the North Equatorial Current (NEC) in the western North Pacific during three consecutive cruises in June and July of 1991 (N=28) and 1994 (N=20), and July and September of 1995 (N=27), indicating that this is a spawning area of this species. These leptocephali were collected from 130°E to 142°E and 12°N to 20°N, to the west of the Mariana Islands, in 20 tows in 1991, in 13 tows in 1994 and in 17 tows in 1995, indicating a widespread presence, but a relatively low abundance. Six of these specimens (16.3-36.0 mm total length) from the 1995 cruise, which were of the typical size range of these leptocephali, were genetically confirmed to be A. marmorata in a previous study. The consistent presence of recently spawned A. marmorata leptocephali (9-20 mm) in all 3 years, suggests that the western region of the NEC is the spawning area of the northern population of A. marmorata that was identified in a recent population genetics study. These leptocephali would thus be transported westward by the NEC and then transported north into the Kuroshio Current and toward Taiwan and Japan, or south toward the southern Philippines and into the Celebes Sea by the Mindanao Current. Available evidence indicates that A. marmorata may have potentially year-round spawning, and the presence of a spawning area of this species in the same region as that of Anguilla japonica suggests that the northern population of A. marmorata has evolved a spawning migration from East Asia, the Philippines and the Celebes Sea region to the NEC area, but differs from A. japonica by having some individuals that recruit to more southern areas.     

Seasonal variation in the degree of heterospecific association of two syntopic hyraxes (<Emphasis Type="Italic">Heterohyrax</Emphasis><Emphasis Type="Italic">brucei </Emphasis>and <Emphasis Type="Italic">Procavia capensis</Emphasis>) exhibiting synchronous parturition

Adult rock hyrax (Procavia capensis) and bush hyrax (Heterohyrax brucei) allocate much of their surface activity to being vigilant for vertebrate predators. Individuals of the two species associate for thermoregulatory basking on rock outcrops (koppies) and, in the Matobo National Park, Zimbabwe, frequently constitute heterospecific groups that produce offspring synchronously. Exposure to predators during basking is thought to put a high premium on vigilance, particularly during the presence of dependent offspring in nurseries. We predicted, therefore, an increase in the association of the two species of hyraxes when offspring were present because this would lead to larger groups, thus enhancing the ability to detect predators. Koppie aggregations of hyraxes were classified as homospecific or heterospecific according to basking group composition. Heterospecific association increased when offspring were present, and heterospecific groups were larger than homospecific ones. Fifty-one percent of nurseries were heterospecific for adults/subadults. During the pre-weaning phase (<2 months after birth), 85% of P. capensis and 65% of H. brucei juveniles were in nurseries of heterospecific basking groups. Heterospecific groups had more offspring than homospecific groups. Juvenile mortality is most evident after young are weaned and heterospecific associations break down. Heterospecific aggregation, through increased group size, may result in a higher probability of avoiding predators, thereby reducing the vulnerability of young.     

Reduced planktotrophy in larvae of <Emphasis Type="Italic">Clypeaster</Emphasis><Emphasis Type="Italic">rosaceus</Emphasis> (Echinodermata,Echiniodea)

Phylogenetic analyses have demonstrated that nonfeeding larvae have evolved from feeding larvae many times among marine invertebrates. In light of this observation, it is surprising that an intermediate strategy, a larva that can feed but is provisioned with enough energy to metamorphose without acquiring exogenous food (i.e., facultative planktotrophy), is rare. A hypothesis for the lack of facultative planktotrophic species among marine invertebrates is that the transition from feeding to nonfeeding is rapid due to this intermediate stage being evolutionarily unstable. Evidence that would support this hypothesis is if species with facultative planktotrophy have reduced food assimilation when compared with obligate planktotrophs. We studied a species with facultative planktotrophic larvae, Clypeaster rosaceus, that is very near the boundary between facultative and obligatory planktotrophy, to answer two questions: (1) does feeding during the larval stage result in energy gains in larval or juvenile stages and (2) if not, are larvae capable of assimilating exogenous food at all. Our measurements of energetics in larval and juvenile stages show that C. rosaceus larvae accumulate very little if any energy when fed, but stable isotope data indicate that larvae are able to assimilate some food. Our results are consistent with similar studies on facultative planktotrophic larvae suggesting poor food assimilation and rapid loss of larval feeding after a population evolves the ability to reach metamorphosis without feeding (lecithotrophy).     

Feeding aggregations of sea stars (<Emphasis Type="Italic">Asterias</Emphasis> spp. and <Emphasis Type="Italic">Henricia sanguinolenta</Emphasis>) associated with sea urchin (<Emphasis Type="Italic">Strongylocentrotus droebachiensis</Emphasis>) grazing fronts in Nova Scotia

Migrating feeding aggregations (or fronts) of sea urchins can dramatically alter subtidal seascapes by destructively grazing macrophytes. While direct effects of urchin fronts on macrophytes (particularly kelps) are well documented, indirect effects on associated fauna are largely unknown. Secondary aggregations of predators and scavengers form around fronts of Strongylocentrotus droebachiensis in Nova Scotia. We recorded mean densities of the sea stars Asterias spp. (mainly A. rubens) and Henricia sanguinolenta of up to 11.6 and 1.7 individuals 0.25 m⁻² along an urchin front over 1 year. For Asterias, mean density at the front was 7 and 15 times greater than in the kelp bed and adjacent barrens, respectively. There was strong concordance between locations of peak density of urchins and sea stars (Asterias r = 0.98; H. sanguinolenta r = 0.97) along transects across the kelp–barrens interface, indicating that sea star aggregations migrated along with the urchin front at rates of up to 2.5 m per month. Size–frequency distributions suggest that Asterias at the front were drawn from both the barrens (smaller individuals) and the kelp bed (larger individuals). These sea stars fed intensively on mussels on kelp holdfasts and in adjacent patches. Urchin grazing may precipitate aggregations of sea stars and other predators or scavengers by incidentally consuming or damaging mussels and other small invertebrates, and thereby releasing a strong odor cue. Consumption of protective holdfasts and turf algae by urchins could facilitate feeding by these consumers, which may obtain a substantial energy subsidy during destructive grazing events.