Genetic divergence in the tribe Electronini (Myctophidae)

To reveal evolutionary relationships and the rate of divergence of lanternfishes in the tribe Electronini (Myctophidae), analysis of sequence of cox 1 mtDNA fragment in three genera (12 species) of the tribe and in a closely related genus Myctophum (9 species) was performed. The results support the tribe monophyly; however, the divergence within it appeared to be not profound: genetic distances between genera and subgenera are smaller than between species within Myctophum and other genera of myctophids. In subgenera of Protomyctophum, species formation has obviously not completed since interspecies distances in them (0.5–4.7%) are comparable to estimates of intraspecific variation in other lanternfishes. Minimal genetic differences were found between species P. (Hierops) arcticum and P. (H.) subparallelum, whose ranges at the present time are isolated in the northern and southern hemispheres. After calibration of the molecular clock, the time of isolation of these two species can be dated to the Middle Pleistocene. Weak tribe diversification indicates its evolutionary youth, which can be related to the absence of efficient hydrographic barriers for isolation. The data obtained do not permit to consider tribe members as the most primitive in the family Myctophidae.     

A new phylogenetic test for comparing multiple high‐dimensional evolutionary rates suggests interplay of evolutionary rates and modularity in lanternfishes (Myctophiformes; Myctophidae)

The interplay between evolutionary rates and modularity influences the evolution of organismal body plans by both promoting and constraining the magnitude and direction of trait response to ecological conditions. However, few studies have examined whether the best‐fit hypothesis of modularity is the same as the shape subset with the greatest difference in evolutionary rate. Here, we develop a new phylogenetic comparative method for comparing evolutionary rates among high‐dimensional traits, and apply this method to analyze body shape evolution in bioluminescent lanternfishes. We frame the study of evolutionary rates and modularity through analysis of three hypotheses derived from the literature on fish development, biomechanics, and bioluminescent communication. We show that a development‐informed partitioning of shape exhibits the greatest evolutionary rate differences among modules, but that a hydrodynamically informed partitioning is the best‐fit modularity hypothesis. Furthermore, we show that bioluminescent lateral photophores evolve at a similar rate as, and are strongly integrated with, body shape in lanternfishes. These results suggest that overlapping life‐history constraints on development and movement define axes of body shape evolution in lanternfishes, and that the positions of their lateral photophore complexes are likely a passive outcome of the interaction of these ecological pressures.     

Oocyte morphology of several mesopelagic fishes in connection with their taxonomic status and habitat conditions

The inner structure of oocytes was studied in 9 species of mesopelagic fish from the Myctophidae, Melamphaidae, Bathylagidae and Platytroctidae families. It was revealed that species of the same family (Myctophidae) differed less in oocyte structure than species from different families. Within the limits of subfamilies the myctophid fishes had the maximum similarity in their oocyte structure. It was established, that oocytes of fishes of different families differ by the size of vacuoles and yolk granules, and by their localization in cytoplasm.     

Morphology of the pineal complex in seven species of lanternfishes (Pisces: Myctophidae)

The morphology of the pineal complex was compared in seven species of lanternfishes (family Myctophidae) using both light and electron microscopes. On the basis of compactness of the pineal end-vesicle and presence or absence of a dorsal sac, the species in this study were divided into two groups. This grouping seems to correlate well with current views on the phylogenetic relationships among these fishes. Receptor cells and supportive cells are described in the pineals of all species examined. The deepest-dwelling of the forms studied, Parvilux ingens, showed a significant increase in the mean number of lamellar membranes in the outer segments of the receptor cells and a higher convergence ratio of receptor cells to ganglion cells as compared to the shallow-dwelling form Tarletonbeania crenularis. Accordingly it is suggested that the pineal of P. ingens is more photosensitive. Additional differences among species were found in the ultrastructure of the supportive cells. Dorsal sacs were absent in the three shallowest-occurring myctophids studied. In those species with a dorsal sac, its close association with the pineal-end-vesicle suggests a functional relationship between the two structures.     

Vision in the deep sea

The deep sea is the largest habitat on earth. Its three great faunal environments--the twilight mesopelagic zone, the dark bathypelagic zone and the vast flat expanses of the benthic habitat--are home to a rich fauna of vertebrates and invertebrates. In the mesopelagic zone (150-1000 m), the down-welling daylight creates an extended scene that becomes increasingly dimmer and bluer with depth. The available daylight also originates increasingly from vertically above, and bioluminescent point-source flashes, well contrasted against the dim background daylight, become increasingly visible. In the bathypelagic zone below 1000 m no daylight remains, and the scene becomes entirely dominated by point-like bioluminescence. This changing nature of visual scenes with depth--from extended source to point source--has had a profound effect on the designs of deep-sea eyes, both optically and neurally, a fact that until recently was not fully appreciated. Recent measurements of the sensitivity and spatial resolution of deep-sea eyes--particularly from the camera eyes of fishes and cephalopods and the compound eyes of crustaceans--reveal that ocular designs are well matched to the nature of the visual scene at any given depth. This match between eye design and visual scene is the subject of this review. The greatest variation in eye design is found in the mesopelagic zone, where dim down-welling daylight and bio-luminescent point sources may be visible simultaneously. Some mesopelagic eyes rely on spatial and temporal summation to increase sensitivity to a dim extended scene, while others sacrifice this sensitivity to localise pinpoints of bright bioluminescence. Yet other eyes have retinal regions separately specialised for each type of light. In the bathypelagic zone, eyes generally get smaller and therefore less sensitive to point sources with increasing depth. In fishes, this insensitivity, combined with surprisingly high spatial resolution, is very well adapted to the detection and localisation of point-source bioluminescence at ecologically meaningful distances. At all depths, the eyes of animals active on and over the nutrient-rich sea floor are generally larger than the eyes of pelagic species. In fishes, the retinal ganglion cells are also frequently arranged in a horizontal visual streak, an adaptation for viewing the wide flat horizon of the sea floor, and all animals living there. These and many other aspects of light and vision in the deep sea are reviewed in support of the following conclusion: it is not only the intensity of light at different depths, but also its distribution in space, which has been a major force in the evolution of deep-sea vision.     

Relationships between otolith size and fish length in some mesopelagic teleosts (Myctophidae,Paralepididae, Phosichthyidae and Stomiidae)

Length–mass relationships and linear regressions are given for otolith size (length and height) and standard length (L_S) of certain mesopelagic fishes (Myctophidae, Paralepididae, Phosichthyidae and Stomiidae) living in the central Mediterranean Sea. The length–mass relationship showed isometric growth in six species, whereas linear regressions of L_S and otolith size fit the data well for all species. These equations represent a useful tool for dietary studies on Mediterranean marine predators.     

Application of geometric morphometrics for intraspecific variability analysis in mesopelagic fishes of Sternoptychidae and Myctophidae families

We present the results of application of geometric morphometrics for studies of variability in populations of the members of two families of South Atlantic mesopelagic fishes, Sternoptyx diaphana (Sternoptychidae) and Ceratoscopelus warmingii (Myctophidae), in which an almost complete genetic isolation was previously discovered within the borders of hydrographic regions. A system of landmark points is developed, describing both anatomical features and configuration of luminescent organs, photophores, which allows to consider not only the influence of external physical and ecological conditions but also the existence of population-specific signaling system as a potential cause of genetic isolation. Morphological similarities are demonstrated for S. diaphana throughout the whole studied portion of the distribution area, while certain differences affecting the head shape and position of individual photophores on the body are discovered between C. warmingii populations.     

Size matters for lice on birds: Coevolutionary allometry of host and parasite body size

Body size is one of the most fundamental characteristics of all organisms. It influences physiology, morphology, behavior, and even interspecific interactions such as those between parasites and their hosts. Host body size influences the magnitude and variability of parasite size according to Harrison's rule (HR: positive relationship between host and parasite body sizes) and Poulin's Increasing Variance Hypothesis (PIVH: positive relationship between host body size and the variability of parasite body size). We analyzed parasite–host body size allometry for 581 species of avian lice (～15% of known diversity) and their hosts. We applied phylogenetic generalized least squares (PGLS) methods to account for phylogenetic nonindependence controlling for host and parasite phylogenies separately and variance heterogeneity. We tested HR and PIVH for the major families of avian lice (Ricinidae, Menoponidae, Philopteridae), and for distinct ecological guilds within Philopteridae. Our data indicate that most families and guilds of avian lice follow both HR and PIVH; however, ricinids did not follow PIVH and the “body lice” guild of philopterid lice did not follow HR or PIVH. We discuss mathematical and ecological factors that may be responsible for these patterns, and we discuss the potential pervasiveness of these relationships among all parasites on Earth.     

Ecological, morphological and phylogenetic correlates of interspecific variation in plasma carotenoid concentration in birds

Carotenoids are important as pigments for bright coloration of animals, and as physiologically active compounds with a wide array of health-related benefits. However, the causes of variation in carotenoid acquisition and physiology among species are poorly known. We measured the concentration of carotenoids in the blood of 80 wild bird species differing in diet, body size and the extent of carotenoid-based traits. Preliminary analyses showed that diet significantly explains interspecific variability in plasma carotenoids. However, dietary influences were apparently overridden by phylogenetic relationships among species, which explained most (65%) of this variability. This phylogenetic effect could be due partly to its covariation with diet, but may also be caused by interspecific differences in carotenoid absorption from food to the blood stream, mediated, for example by endothelial carriers or gut parasites. Carotenoid concentrations also decreased with body size (which may be explained by the allometric relationship between ingestion rate and body mass), and correlated positively with the extent of carotenoid-dependent coloration of plumage and bare parts. Therefore, the acquisition of carotenoids from the diet and their use for both health and display functions seem to be constrained by ecological and physiological aspects linked to the phylogeny and size of the species.     

Feeding ecology of Mediterranean common dolphins: The importance of mesopelagic fish in the diet of an endangered subpopulation

The Mediterranean subpopulation of common dolphin (Delphinus delphis) is classified as endangered by the IUCN. Still, information about their diet in the Mediterranean is scarce. Stomach contents of 37 common dolphins stranded in the Alboran Sea and Strait of Gibraltar were analyzed. A total of 13,634 individual prey of 28 different taxa were identified. For fish, Myctophidae was the most important family as indicated by the highest index of relative importance (IRI = 8,470), followed by the family Sparidae (IRI = 609). The most important Myctophidae species was Madeira lantern fish (Ceratoscopelus maderensis) and for Sparids, the bogue (Boops boops). Cephalopods, instead, were found in low quantities only with 31 prey from the Loliginidae, Ommastrephidae, and Sepiolidae families. Overall, our results indicate that common dolphins are mainly piscivorous (99.77%N, 94.59%O, 99.73%W), feeding mostly on mesopelagic prey. Although common dolphins inhabit mainly coastal waters in the study area, the narrow continental shelf seems to facilitate the availability of Myctophids and other members of the mesopelagic assemblage to dolphins when the assemblage migrates to the surface at night. Our results represent the first attempt at quantifying the diet of this predator in the Alboran Sea and Strait of Gibraltar.     

The eyes of deep-sea fishes and the changing nature of visual scenes with depth

The visual scenes viewed by ocean animals change dramatically with depth. In the brighter epipelagic depths, daylight provides an extended field of illumination. In mesopelagic depths down to 1000 m the visual scene is semi-extended, with the downwelling daylight providing increasingly dim extended illumination with depth. In contrast, greater depths increase the prominence of point-source bioluminescent flashes. In bathypelagic depths (below 1000 m daylight no longer penetrates, and the visual scene consists exclusively of point-source bioluminescent flashes. In this paper, I show that the eyes of fishes match this change from extended to point-source illumination, becoming increasingly foveate and spatially acute with increasing depth. A sharp fovea is optimal for localizing point sources. Quite contrary to their reputation as 'degenerate' and 'regressed', I show here that the remarkably prominent foveae and relatively large pupils of bathypelagic fishes give them excellent perception and localization of bioluminescent flashes up to a few tens of metres distant. In a world with almost no food, where fishes are weak and must swim very slowly this range of detection (and interception) is energetically realistic, with distances greater than this physically beyond range. Larger and more sensitive eyes would give bathypelagic fishes little more than the useless ability to see flashes beyond reach.     

What determines host range in parasitoids? An analysis of a tachinid parasitoid community

Despite the vast diversity of parasitic insects and their importance in natural and agricultural communities, our knowledge of what determines their patterns of association with hosts remains sparse. Unlike most parasites that tend to be specialized, parasitoid flies in the family Tachinidae exhibit a broad spectrum of host-specificity, with many species attacking a wide range of hosts. This variability in host-specificity makes them a useful model for examining the ecological and historical factors that determine host associations. We analyzed data collected from a 5-year rearing program of Lepidoptera in southern Arizona to investigate the factors that influence tachinid-host associations. After controlling for a strong effect of sample size, a significant portion of the remaining variance in host range was explained by differences among phylogenetic groups of tachinids and/or their correlated reproductive strategies. Relatively specialized tachinids tended to be associated with monophagous or narrowly oligophagous hosts and attacked them at relatively high frequencies, a pattern we suggest is related to host location efficiency. Cluster analysis indicated that host abundance, gregariousness, food-plant type, and morphology are all important determinants of tachinid host use. Little concordance was found between how tachinid species cluster according to characteristics of their hosts and their estimated phylogenetic relationships. Together, the results of this study suggest that ecological factors are important determinants of host use in these parasitoids and although phylogenetic history may influence the range of hosts used, its power to explain the ecological or taxonomic character of hosts used appears limited.     

Ecology,song similarity and phylogeny predict natural hybridization in an avian family

Hybridization is important in the evolution of many animal groups; however, broad scale patterns of natural hybridization are still poorly understood. Using phylogenetic comparative analyses, we tested for relationships between demographic, ecological, phenotypic and phylogenetic variables and the incidence of natural hybridization among 45 species of North American wood warbler. Since 1980, hybrids have been documented in 24 species (53 %). We detected negative relationships between the incidence of hybridization and both breeding range size and phylogenetic distance, and positive relationships between the incidence of hybridization and (1) song similarity, (2) the extent of breeding sympatry of species pairs and (3) the number of additional heterospecific warbler species that co-occur during breeding. Neither population size nor breeding habitat quality (as measured by threats to survival or reproduction in breeding areas) explained variation in the incidence of hybridization. Our results suggest a potential role of limited breeding habitat in wood warbler hybridization events and a net positive effect of breeding sympatry. They also support the hypothesis that mating signal similarity facilitates hybridization events and are consistent with an increase in reproductive isolation with increasing genetic divergence. Our results also suggest the presence of phylogenetic signal in wood warbler hybridization. By investigating natural hybridization at the taxonomic level of family, we have identified several previously undocumented patterns of natural hybridization. This study demonstrates the utility of examining the combined effects of ecology, demography, phenotype and phylogeny when studying variation in the expression of natural hybridization among taxa.     

Comparative study of the spectral sensitivities of mesopelagic crustaceans

The spectral sensitivities of 12 species of mesopelagic crustaceans were studied by means of electrophysiological recordings. Nine of the species are vertical migrators, while 3 are not, and 9 species possess bioluminescent organs, while 3 are not bioluminescent. All species had a single peak of spectral sensitivity with maxima between 470 nm and 500 nm. There was no apparent correlation between sensitivity maxima and daytime depth distribution, migratory behavior, or the presence or absence of bioluminescent organs. With the exception of the hyperiid amphipod Phronima sedentaria, the spectral sensitivities of these mesopelagic crustaceans demonstrate a better match for maximum sensitivity to bioluminescence than to downwelling light. Accepted: 29 June 1999     

Phylogenetic resolution and habitat specificity of members of the Photobacterium phosphoreum species group

Substantial ambiguity exists regarding the phylogenetic status of facultatively psychrophilic luminous bacteria identified as Photobacterium phosphoreum, a species thought to be widely distributed in the world's oceans and believed to be the specific bioluminescent light-organ symbiont of several deep-sea fishes. Members of the P. phosphoreum species group include luminous and non-luminous strains identified phenotypically from a variety of different habitats as well as phylogenetically defined lineages that appear to be evolutionarily distinct. To resolve this ambiguity and to begin developing a meaningful knowledge of the geographic distributions, habitats and symbiotic relationships of bacteria in the P. phosphoreum species group, we carried out a multilocus, fine-scale phylogenetic analysis based on sequences of the 16S rRNA, gyrB and luxABFE genes of many newly isolated luminous strains from symbiotic and saprophytic habitats, together with previously isolated luminous and non-luminous strains identified as P. phosphoreum from these and other habitats. Parsimony analysis unambiguously resolved three evolutionarily distinct clades, phosphoreum, iliopiscarium and kishitanii. The tight phylogenetic clustering within these clades and the distinct separation between them indicates they are different species, P. phosphoreum, Photobacterium iliopiscarium and the newly recognized 'Photobacterium kishitanii'. Previously reported non-luminous strains, which had been identified phenotypically as P. phosphoreum, resolved unambiguously as P. iliopiscarium, and all examined deep-sea fishes (specimens of families Chlorophthalmidae, Macrouridae, Moridae, Trachichthyidae and Acropomatidae) were found to harbour 'P. kishitanii', not P. phosphoreum, in their light organs. This resolution revealed also that 'P. kishitanii' is cosmopolitan in its geographic distribution. Furthermore, the lack of phylogenetic variation within 'P. kishitanii' indicates that this facultatively symbiotic bacterium is not cospeciating with its phylogenetically divergent host fishes. The results of this fine-scale phylogenetic analysis support the emerging view that bacterial species names should designate singular historical entities, i.e. discrete lineages diagnosed by a significant divergence of shared derived nucleotide characters.     

Species distribution,ecology, abundance,body size and phylogeny originate interrelated rarity patterns at regional scale

The most pervasive macroecological patterns concern (1) the frequency distribution of range size, (2) the relationship between range size and species abundance and (3) the effect of body size on range size. We investigated these patterns at a regional scale using the tenebrionid beetles of Latium (Central Italy). For this, we calculated geographical range size (no. of 10‐km square cells), ecological tolerance (no. of phytoclimatic units) and abundance (no. of sampled individuals) using a large database containing 3561 georeferenced records for 84 native species. For each species, we also calculated body mass and its ‘phylogenetic diversity’ on the basis of cladistic relationships. Frequency distribution of range size followed a log‐normal distribution as found in many other animal groups. However, a log‐normal distribution accommodated well the frequency distribution of ecological tolerance, a so far unexplored issue. Range size was correlated with abundance and ecological tolerance, thus supporting the hypothesis that a positive correlation between distribution and abundance is a reflection of interspecific differences in ecological specialization. Larger species tended to have larger ranges and broader ecological tolerance. However, contrary to what known in most vertebrates, not only small‐sized, but also many medium‐to‐large‐sized species exhibited great variability in their range size, probably because tenebrionids are not so strictly influenced by body size constraints (e.g. home ranges) as vertebrates. Moreover, in contrast to other animals, tenebrionid body size does not influence species abundances, probably because these detritivorous animals are not strongly regulated by competition. Finally, contrary to the assumption that rare species should be mainly found among lineages that split from basal nodes, rarity of a tenebrionid species was not influenced by the phylogenetic position of its tribe. However, lineages that split from more basal nodes had lower variability in terms of species geographical distribution, ecological tolerance and abundance, which suggests that lineages that split from more basal nodes are not only morphologically conservative but also tend to have an ecological ‘inertia’.     

New observations and ontogenetic transformation of photogenic tissues in the tubeshoulder Sagamichthys schnakenbecki (Platytroctidae,Alepocephaliformes)

Several species of the luminescent tubeshoulder fish (family Platytroctidae) show extensive ontogenetic transformations in the development of bioluminescent structures from larvae to adults. Several types of luminescent tissues are present in platytroctids, although these tissues are poorly known for most species because specimens are rarely observed. The present study describes the ontogenetic transformation of photogenic structures in Sagamichthys schnakenbecki, a species that is found in meso and bathy-pelagic depths of the Atlantic Ocean. Five newly described luminous structures are included in addition to a review of all known bioluminescent tissues described in the family. The newly discovered photogenic tissues were observed at the pectoral-fin base in early juveniles, as a pair of large globule-like tissues inside the caudal peduncle of early juveniles, at the pelvic girdle of late juveniles and early adults and as photogenic tissue observed as pigment over the cleithral bone in adults. A peculiar skin-slit structure, which was observed only in S. schnakenbecki, is described and discussed. Skin slits were associated with certain bioluminescent structures during the transformation into adulthood. In addition, coI sequence data from nine of 13 recognized platytroctid genera were used to construct the first molecular phylogenetic tree for the family. Finally, the first photographic evidence of the rarely observed luminous discharge of a tubeshoulder shoulder organ is presented from observations off south-east Greenland.     

Photon Hunting in the Twilight Zone: Visual Features of Mesopelagic Bioluminescent Sharks

The mesopelagic zone is a visual scene continuum in which organisms have developed various strategies to optimize photon capture. Here, we used light microscopy, stereology-assisted retinal topographic mapping, spectrophotometry and microspectrophotometry to investigate the visual ecology of deep-sea bioluminescent sharks [four etmopterid species (Etmopterus lucifer, E. splendidus, E. spinax and Trigonognathus kabeyai) and one dalatiid species (Squaliolus aliae)]. We highlighted a novel structure, a translucent area present in the upper eye orbit of Etmopteridae, which might be part of a reference system for counterillumination adjustment or acts as a spectral filter for camouflage breaking, as well as several ocular specialisations such as aphakic gaps and semicircular tapeta previously unknown in elasmobranchs. All species showed pure rod hexagonal mosaics with a high topographic diversity. Retinal specialisations, formed by shallow cell density gradients, may aid in prey detection and reflect lifestyle differences; pelagic species display areae centrales while benthopelagic and benthic species display wide and narrow horizontal streaks, respectively. One species (E. lucifer) displays two areae within its horizontal streak that likely allows detection of conspecifics'' elongated bioluminescent flank markings. Ganglion cell topography reveals less variation with all species showing a temporal area for acute frontal binocular vision. This area is dorsally extended in T. kabeyai, allowing this species to adjust the strike of its peculiar jaws in the ventro-frontal visual field. Etmopterus lucifer showed an additional nasal area matching a high rod density area. Peak spectral sensitivities of the rod visual pigments (λ_max) fall within the range 484–491 nm, allowing these sharks to detect a high proportion of photons present in their habitat. Comparisons with previously published data reveal ocular differences between bioluminescent and non-bioluminescent deep-sea sharks. In particular, bioluminescent sharks possess higher rod densities, which might provide them with improved temporal resolution particularly useful for bioluminescent communication during social interactions.     

Insight into an island radiation: the Tarentola geckos of the Cape Verde archipelago

Aim To reassess the relationships between Tarentola geckos from the Cape Verde Islands by including specimens from all islands in the range. To determine the variation within forms by sequencing over 400 specimens, thereby allowing the discovery of cryptic forms and resolving some of the issues raised previously. This extensive sampling was also used to shed light on distributions and to explain genetic diversity by comparing the ages and ecological and geological features of the islands (size, elevation and habitat diversity). Location The Cape Verde Islands: an oceanic archipelago belonging to the Macaronesian biogeographic region, located around 500 km off Senegal. Methods A total of 405 new specimens of Tarentola geckos were collected from nine islands with very different geological histories, topography, climate and habitats. Mitochondrial cytochrome b (cyt b) gene and 12S rRNA partial sequences were obtained and analysed using phylogenetic methods and networks to determine molecular diversity, demographic features and phylogeographic patterns. Results The phylogenetic relationships between all known forms of Cape Verdean Tarentola specimens were estimated for the first time, the relationships between new forms were assessed and previously hypothesized relationships were re‐examined. Despite the large sample size, low intraspecific diversity was found using a 303‐bp cyt b fragment. Star‐like haplotype networks and statistical tests suggest the past occurrence of a rapid demographic and geographical expansion over most of the islands. Genetic variability is positively correlated with size, elevation and habitat diversity of the islands, but is not linearly related to the age of the islands. Biogeographical patterns have, in general, high concordance with phylogenetic breaks and with the three eco‐geographical island groups. Volcanism and habitat diversity, both tightly linked with island ontogeny, as postulated by the general dynamic model of oceanic island biogeography, as well as present and historical size of the islands appear to be the main factors explaining the genetic diversity of this group. Main conclusions The Tarentola radiation was clarified and is clearly associated with the geological and ecological features of the islands. Two factors may account for the low intraspecific variation: (1) recent volcanic activity and high ecological stress, and (2) poor habitat diversity within some islands. More studies are needed to align taxonomy with phylogenetic relationships, whereas GIS modelling may help to predict precise species distributions.     

Diet of the King Penguin Aptenodytes patagonicus during three summers at South Georgia

King Penguins Aptenodytes patagonicus which were rearing chicks were studied during three summers from November 1991 to March 1994 at South Georgia. Stomach samples ( n =115) collected by flushing had a mean mass of 1308 g. Fish mass was allocated to each species based on the relationship between fish mass and otolith length. Three mesopelagic lanternfishes (Myctophidae), Krefftichthys anderssoni, Electrona carlsbergi and Protomyctophum choriodon , dominated the diet both by numbers and mass. They were small fish with mean mass of 3–7 g. Overall, K. anderssoni dominated the diet in terms of numbers and mass. Although Barracudina Notolepis coatsi occurred in <3% of the diet by numbers, it was large (106 g) and was second most important in terms of mass. Squid represented <3% of the diet by mass. Although the chick-rearing success was poor in the 1993–1994 summer, meal size was not reduced but foraging trips were longer. In the 1993–1994 summer, a larger proportion of the otoliths were not identifiable because they were more completely digested. Fewer otoliths were identified as being those of K. anderssoni , but we argue that about 90% of the unidentified otoliths were K. anderssoni. There was also more squid and N. coatsi in the diet during the poor summer. A consistent trend was that P. choriodon was rare or absent in early summer but more important later in the year, and at the end of 1992–1993, it was the dominant prey. We conclude that myctophid fish, especially K. anderssoni , are the main summer prey of King Penguins rearing chicks at South Georgia, as found in other recent studies in the Southern Ocean.