Seasonal changes in the diving parameters of king penguins (Aptenodytes patagonicus)

Contrasting conditions at-sea are likely to affect the foraging behaviour of seabirds. However, the effect of season on the dive parameters of penguins is poorly known. We report here on an extensive study of the diving behaviour of king penguins (Aptenodytes patagonicus) over the bird's complete annual cycle at the Crozet Islands. Time-depth recorders were used to record dive duration, bottom duration, post-dive interval, ascent rate and descent rate in breeding adults during different seasons in 1995 and 1996. Seasons included summer (n=6, incubation; n=6, chick brooding), autumn and winter (n=5 and n=3, respectively, chick at the crèche stage), and spring (n=4, birds at the post-moult stage). In all seasons dive duration increased with dive depth, but, for a given depth, dives were longer in winter (6.8 min when averaged over the 100-210 m depth layer) than in spring (4.6 min) and summer (4.4 min). The time spent at the bottom of the dives, which probably represents a substantial part of the feeding time, was much longer in winter (2.5 min per dive for dives over the 100-210 m layer) than during other seasons (1.0-1.4 min), i.e. there was a 2.5-fold augmentation for similar diving depths. Ascent and descent rates increased with increasing dive depth, but no difference in the relationships between rates of ascent and descent and dive depth was found among seasons. Furthermore, for all dive depths, ascent and descent rates were independent of the bottom duration. In all seasons post-dive intervals increased with dive duration and with dive depth, but they were longer in spring (2.3 min for dives over the 100-210 m layer) and summer than in autumn and winter (1.6-1.8 min). The diving efficiency decreased with increasing dive depth and was higher in autumn and winter (0.22-0.29) than in summer and spring (0.15-0.18). The large increase in bottom and dive duration from spring to winter is in agreement with the seasonal drop in prey density, with penguins spending more time searching for prey. In contrast, the consistency of the vertical velocity during contrasting conditions at-sea suggests that the transit time to depth is an important component of the foraging behaviour (scanning of the water column) that is independent of the prey availability. The time budget of the penguins during diving in a fluctuating environment appears to vary primarily during the bottom phase of the dives, with bottom duration increasing with diminishing prey supplies, while post-dive intervals shorten in the same time.     

Fish and squid in the diet of king penguin chicks,Aptenodytes patagonicus,during winter at sub-antarctic Crozet Islands

The diet of king penguins, Aptenodytes patagonicus, rearing chicks was studied during three consecutive austral winters (1990, 1991 and 1992) at Crozet Islands. The mean stomach content mass of the 47 samples was 503 g. Percentages of wet and reconstituted masses showed that both fishes (66 and 36%, respectively) and squid (34 and 64%) are important components of the winter diet. Juveniles of the demersal onychoteuthid squid Moroteuthis ingens form the bulk of the cephalopod diet, and this was the main prey by reconstituted mass (57%). Myctophid fish (lantern-fishes) accounted for most of the fish diet, constituting together 32% by mass. The three main species of myctophids eaten in summer by king penguins were either very rare in winter (Electrona carlsbergi) or accounted for a smaller proportion of the diet (Krefftichthys anderssoni = 1.5% by mass and Protomyctophum tenisoni = 4.6%). Five other myctophids, which are rarely consumed in summer, contributed 24% of the diet by mass in winter (Gymnoscopelus piabilis = 18.1%, Lampichthys procerus = 2.4%, G. nicholsi = 1.3%, and Metelectrona ventralis and Electrona subaspera = 1.0%). The greater diversity of prey in winter suggests a more opportunistic feeding behaviour at a time probably marked by a change in prey availability. Both the known ecology of the fish and squid prey and the barely digested state of some items suggest that in winter breeding adults forage in the outer shelf, upper slope and oceanic areas in the close vicinity of the Crozet Islands to feed their chicks. Finally, using king penguins as biological samplers, the present work provides novel data on the previously unstudied mesopelagic/epibenthic marine community in waters surrounding the Crozet Islands. Seventeen myctophid fish have been identified to species level. These include several poorly known species in the southern Indian Ocean. The occurrence of small, nearly intact, cephalopods in the diet of king penguins suggests that spawning grounds of four squid species may be located near the Crozet Archipelago.     

Fatty acid signature analysis documents the diet of five myctophid fish from the Southern Ocean

Fatty acid (FA) and fatty alcohol (FAlc) compositions of both total lipid and neutral lipid fractions were studied for five myctophid species sampled in Kerguelen waters. Both qualitative and quantitative FA signature analyses were then performed to investigate their diet over longer time scales than the conventional stomach content analysis. Regarding their lipid class, FA and FAlc compositions, the five species could be discriminated into two groups: wax-ester-rich species (Electrona antarctica, Krefftichthys anderssoni) characterised by large amounts of monounsaturated FAs (>73% of total FAs) and triacylglycerol-rich species (Electrona carlsbergi, Gymnoscopelus nicholsi, Protomyctophum bolini) with major amounts of saturated and monounsaturated FAs (>29 and >46% of total FAs, respectively). Qualitative and quantitative FA analyses showed that K. anderssoni mainly preyed upon copepods, E. antarctica upon copepods and more euphausiids and P. bolini and E. carlsbergi mainly upon euphausiids with some copepods, while G. nicholsi had a more diverse diet. This study shows the usefulness of quantitative statistical analysis to determine the diet of Antarctic and sub-Antarctic predators and stresses the need of increasing the lipid and FA analyses of more zooplanktonic and micro-nektonic marine species.     

Comparison of feeding habits of myctophid fishes and juvenile small epipelagic fishes in the western North Pacific

To examine the potential trophic competition between myctophids and small epipelagic fishes in the nursery grounds in spring, we compared the stomach contents of dominant myctophids (Symbolophorus californiensis, Ceratoscopelus warmingii and Myctophum asperum; n = 179) and juvenile epipelagic fishes (Japanese sardine, Sardinops melanostictus, Japanese anchovy, Engraulis japonicus, chub mackerel, Scomber japonicus, and spotted mackerel, S. australasicus; n = 78) that were simultaneously collected at nighttime with a midwater trawl net around the Kuroshio-Oyashio transition zone in the western North Pacific. It was clear that the neritic copepod Paracalanus parvus s.l. was the most abundant species in NORPAC samples (0.335 mm mesh size) taken at the same stations. Diets of dominant myctophid fishes differed from those of the juvenile epipelagic fishes; Japanese sardine and anchovy mostly preyed upon P. parvus s.l. (23.6% of stomach contents in volume) and Corycaeus affinis (16.1%), respectively. Both chub and spotted mackerels mainly preyed upon the seasonal vertical migrant copepod, Neocalanus cristatus (15.9 and 14.7%, respectively). On the contrary, myctophid fishes probably do not specifically select the abundant neritic copepods. Namely, S. californiensis mostly preyed upon a diel vertical migrating copepod, Pleuromamma piseki (22.7 and 30.6% in stomach of juvenile and adult, respectively), while C. warmingii and M. asperum preyed on Doliolida (43.0% in stomach of juvenile C. warmingii), appendicularians (11.0% in stomach of juvenile M. asperum), and Ostracoda (6.3% in stomach of adult C. warmingii). Feeding habits of myctophid fishes seem adapted to their prey animals; low rate of digested material (less than 30% in volume) in stomachs of S. californiensis may be linked to the movement of P. piseki, hence S. californiensis can easily consume this copepod at night since they are more concentrated at night than daytime. High rate of digested material (over 40%) of M. asperum and adult C. warmingii suggest that they feed not only at night but also during the daytime in the midwater layer. Thus, myctophid fishes actually fed in the surface layer but less actively than the small epipelagic fishes. These results suggest that the potential for direct food competition between myctophids and small epipelagic fishes is low in the nursery ground, but there remains a possibility of indirect effects through their prey items, since the above gelatinous animals feed on common prey items as juveniles of Japanese sardine and anchovy.     

Distribution patterns of larval myctophid fishes in the transition region of the western North Pacific

We examined the species composition and horizontal distribution of myctophid fish larvae in the transition region of the western North Pacific during the early summer. In total, 4,760 myctophid larvae were collected at 44 stations; 18 species of myctophids from 15 genera were collected, and the 8 most abundant species accounted for >95% of larvae. The distribution patterns of these larvae were well defined by the hydrographic structures of the study area including the Oyashio and Kuroshio fronts, the Subarctic Boundary, and a warm core ring. The horizontal distribution patterns of the eight dominant species were categorized into three groups: northern transition water (Stenobrachius nannochir, Tarletonbeania taylori, and Lampanyctus jordani), southern transition water (Symbolophorus californiensis, Diaphus theta, and Nannobrachium regale), and Kuroshio (Myctophum asperum and Diaphus garmani). The Subarctic Boundary defined the distributions of the northern and southern transition-water groups. The importance of areas of western North Pacific transition water as spawning and nursery grounds for subarctic, transitional, and subtropical myctophid fishes was indicated by the relationship between the horizontal distribution patterns of larvae, juveniles, and adults and the physical oceanographic structures.Communicated by T. Ikeda, Hakodate     

Observations on the comparative morphology and operation of the photogenic tissues of myctophid fishes

Lanternfishes (Myctophidae) often bear not only small primary photophores, but also larger regions of luminous tissue, described as glands or patches. These have a superficial appearance very different to that of the primary photophores, but examination of the tissues of some 14 genera has shown that all the luminous organs are composed of fundamentally similar stacks of photogenic lamellae. The degree of organisation of the lamellae varies in different genera, but the caudal organs and the other larger luminous regions contain much more of this photogenic tissue than do the primary photophores. Some observations on the operation of both primary and caudal organs are described.     

The diving behaviour of brooding king penguins (<Emphasis Type="Italic">Aptenodytes patagonicus</Emphasis>) from the Falkland Islands: variation in dive profiles and synchronous underwater swimming provide new insights into their foraging strategies

The diving behaviour of king penguins (Aptenodytes patagonicus) was studied on the Falkland Islands, where a small population (ca. 300 fledglings year^–1) is located at the geographical limit of their breeding range. King penguins rearing newly hatched chicks were equipped with time-depth recorders before leaving for sea. In total, 20,175 dives >3 m were recorded from 12 birds during 15 foraging trips with a mean duration of 5.7±2.3 days. The majority of the trips was directed up to 500 km to the northeast of the breeding colony in slope waters of, and oceanic waters beyond, the Patagonian shelf. Mean time spent underwater accounted for 42±9% of the foraging trip. Mean dive depth achieved was 55±16 m; maximum dive depth recorded was 343 m. Mean dive duration was 159±25 s; maximum dive duration was 480 s. The mean vertical distance covered was 140±65 km trip^–1; and on average birds covered 25 km day^–1. Synchronous diving behaviour was observed in two birds for a period of about 24 h after leaving the colony. Dive depth correlated positively with: (1) light intensity, (2) dive duration and (3) vertical velocities, thus confirming previous findings obtained from conspecifics at other breeding sites and indicating comparable diving behaviour. However, separation of dives according to their profile—V-, U-, or W-shaped—revealed significant differences between certain dive parameters. For a given depth range, bottom time was longer and vertical velocities higher in W-dives than in U-dives. This, together with a higher number of W-dives at dawn and dusk, suggests that foraging is more effective during W-dives than U-dives, and during twilight. These findings imply that king penguins have to make more complex decisions, individually and socially, on the performance of the subsequent dive than previously thought.Communicated by O. Kinne, Oldendorf/Luhe     

Spawning behaviour,egg and larval development of the myctophid fish Benthosema pterotum

The distribution of eggs and early larval stages of the myctophid fish Benthosema pterotum was studied during three cruises to the Gulf of Oman during January to February 1981 and February 1983. The eggs and yolk sac larvae were identified and described after artificial fertilization on board the Research Vessel Dr. Fridtjof Nansen. The mature fish spawn in the Gulf at 300 to 100 m depths during early night. The eggs and smallest larvae were found at 300 to 200 m depths. The eggs hatch within 12 h at 21°C. Hatching occurs before the slightly bouyant eggs reach the upper 50 m of the water column.     

Positive indirect effects of reef fishes on kelp performance: the importance of mesograzers

It has been suggested that microcarnivorous reef fishes may play an important role in giant kelp forest communities by preventing infestations of mesograzers that could severely impact or potentially destroy recovering kelp forests after extreme disturbance events. However, these trophic linkages, specifically the direct and indirect effects of fishes on the biomass of mesograzers, grazing intensity, and the performance of giant kelp, have not been sufficiently quantified and evaluated as to their importance and in the absence of such disturbance events. We examined experimentally the effects of mesograzers on the growth and performance of giant kelp in the presence and absence of their fish predators near Santa Catalina Island, California (U.S.A.). Mesograzer biomass and grazing intensity were significantly higher when fishes were excluded from giant kelp, which in turn, lowered kelp performance. This pattern was consistent both on experimental plots of kelp as habitat isolates, and on a continuous reef. Moreover, the abundance of mesograzers was inversely related to the abundance of kelp perch among several kelp-forested reefs, suggesting that these effects can occur at larger spatial scales. Because of differences in the diet and behavior of two microcarnivorous fishes, the kelp perch and se?orita, we conducted an experiment manipulating each species and its density independently to determine their separate effects on mesograzers and kelp performance. Concurrently we examined the growth and mortality of juvenile kelp. Grazing intensity decreased, estimates of kelp performance increased, and the growth of juvenile kelp increased with increasing densities of fish but with no detectable effects between fishes. Our results demonstrate that these microcarnivorous fishes have positive indirect effects on kelp performance by reducing mesograzer biomass and grazing intensity, and the early life stages of other fishes also may be important. More specifically, these fishes have a positive effect on the density of fronds of giant kelp that can result in greater recruitment success and the abundance of kelp-associated invertebrates and fishes. Indeed, this study suggests that mesograzers have the potential to be one of the most important herbivores in kelp forest ecosystems.     

Topological analysis of the ecological importance of elasmobranch fishes: A food web study on the Gulf of Tortugas, Colombia

We built a trophic network based on a matrix of interspecific trophic relationships to assess the role of elasmobranch fishes in shaping community structure of the Gulf of Tortugas in the Colombian Pacific Ocean. We analyzed diet similarities to define trophic components (nodes) - rather than taxonomical groups - in the network. We evaluated the ecological function of species or trophic entities through topological analysis of their structural importance in trophic networks by applying one local and several mesoscale network indices. We found that top predatory elasmobranchs play an important ecological role in top-down control and in propagating indirect effects through the system owing to high values of the node degree, centrality and topological importance indices. However, invertebrates and teleost fishes had higher connectivity and topological importance than other elasmobranchs in the network before and after removal of top predators from the system. Results from our study thus suggest that elasmobranchs at intermediate trophic levels - commonly referred to as “mesopredators” - are not so important in all complex coastal ecosystems as previously reported.     

Seasonal changes in the diet of a critically endangered seabird and the importance of trawling discards

Pelagic seabirds obtain food from oceans where the availability of their prey changes rapidly both seasonally and spatially. Here, we investigated changes in the trophic habits of the critically endangered Balearic shearwater (Puffinus mauretanicus) through the breeding season and tested for dietary differences between sexes and age classes. We analysed δ¹⁵N and δ¹³C values in blood of adults during the pre-incubation, incubation and chick-rearing periods and of their chicks. Using a two-isotope mixing model, we estimated dietary contributions based on isotope values from potential prey species which included small pelagic species available naturally and demersal fish species available only from trawling discards. Balearic shearwaters showed clear isotopic and dietary variation through the breeding season. During pre-incubation, breeding adults appeared to exploit demersal fish, whereas during the incubation and chick-rearing period, they fed mainly on pelagic anchovies (Engraulis encrasicolus) and pilchards (Sardina pilchardus). Similarly, chicks were fed mainly with anchovies, a resource with a high energetic value. This variation in the dietary habits of adult shearwaters during the breeding season was probably related to both natural and fishery-induced seasonal changes in the availability of potential prey species within their main feeding grounds. However, changes in the nutritional requirements of the shearwaters could also play an important role. Indeed, diet differed between sexes during pre-incubation: females fed less on trawling discards and more on small pelagic fish than males. This sexual segregation in diet could be the consequence of higher nutritional requirements of females during this period. Our study reveals the differential importance of both trawling discards and small pelagic fish species for a pelagic seabird depending on the breeding period and illustrates the importance of considering the entire breeding season when making inferences about the importance of specific prey in seabird dietary studies.     

Metabolic cold adaptation in Antarctic fishes: evidence from enzymatic activities of brain

To evaluate the concept of metabolic cold adaptation (MCA) in fishes, we compared - in brain, red muscle, and white muscle of Antarctic notothenioid fishes and tropical/subtropical fishes - the activities of two enzymes of ATP-generating pathways, citrate synthase (CS), an indicator of citric acid cycle activity (aerobic metabolism), and lactate dehydrogenase (LDH), an indicator of potential for ATP production through anaerobic glycolysis. Brain was chosen because, unlike locomotory muscle, its metabolic activity is not likely to be influenced by a species' level of activity or nutritional status, so MCA should be readily observed if present. CS and LDH activities in brain exhibited a high level of MCA, but compensation to temperature was not complete (48% for CS; 46% for LDH). CS and LDH activities in red and white muscle varied widely among species, according to the general level of locomotory activity. The 'mode of life'-related enzymatic activities in locomotory muscle show that study of MCA at the level of whole organism metabolism is fraught with difficulties and experimental ambiguities. In contrast, the low variation among species within each group in enzymatic activities in brain, and the large differences between groups in CS and LDH activity, show that brain is an excellent study system for evaluating metabolic compensation to temperature.     

Digestive enzymes in the ontogenetic stages of the southern king crab, Lithodes santolla

The early ontogenetic stages of the sub-Antarctic king crab Lithodes santolla were analyzed for the presence and activities of a set of important digestive enzymes. The eggs and non-feeding larvae (zoea I-III, megalopa) showed high activities of esterases, phosphatases, and exopeptidases indicating the enzymatic ability to utilize endogeneous yolk reserves. SDS-PAGE showed a continuous decrease of proteins or proteids in the range of 59–81 kDa during ontogenetic development from the eggs through the zoeal stages to the first juvenile crab stage, CI. This reduction reflects the degradation of storage compounds during lecithotrophic larval development. Activities of the endopeptidases, trypsin and chymotrypsin, were low in eggs and larvae but increased significantly in the first juvenile crab stage. These enzymes typically facilitate the first steps of proteolysis in the extra-cellular spaces of the midgut gland and in the stomach. Their scarcity indicates that the larvae of L. santolla are physiologically not prepared to digest external food. This ability seems to appear first in the CI stage. Extracts of juvenile midgut glands and the gastric fluids of adults showed high activities of a variety of digestive enzymes including phosphatases, carbohydrases, as well as endo- and exopeptidases. High activities of digestive enzymes in adults may compensate for scarce food supply and rate-limiting low temperatures in the predominantly sub-Antarctic habitats of L. santolla.     

Short- and long-term importance of small sharks in the diet of the rare deep-sea shark Dalatias licha

Knowing the trophic ecology of marine predators is essential to develop an understanding of their ecological role in ecosystems. Research conducted on deep-sea and threatened shark species is limited. Here, by combining analyses of individual stomach contents and stable isotope values, we examined the trophic ecology (dietary composition and trophic position) of the kitefin shark Dalatias licha, a deep-sea shark considered as near threatened globally and as data deficient in the Mediterranean Sea. Results revealed the importance of small sharks in the diet of the kitefin shark at short- and long-term scales, although fin-fish, crustaceans and cephalopods were also found. Predation on sharks reveals the high trophic position of the kitefin shark within the food web of the western Mediterranean Sea. Stable isotope values from liver and muscle tissues confirmed our results from stomach content analysis and the high trophic position.     

Evaluating the relative importance of intrinsic and extrinsic factors on the foraging activity of top predators: a case study on female little penguins

A complex interaction of biotic and abiotic factors influences animal foraging activity. It is often difficult to understand which factors may affect animals’ foraging and how it is affected. For instance, whereas the effect of sexual dimorphism on foraging activity has been reported in several species, little is known of the complex interactions between variables acting at a finer scale, e.g. the variability of body mass within a sex. Evaluating the importance of these finer scale factors is also essential to the understanding of foraging behaviour. We propose here a simple approach by applying principal component analysis (PCA) in a novel way to examine relationships between biotic and abiotic factors affecting foraging behaviour of top predators. We studied female little penguins (Eudyptula minor) of known age, carrying miniature accelerometers during the guard stage of breeding. Surprisingly, the body mass of the females did not influence any of the foraging parameters, but females foraging later in the breeding season dived shallower and more often, showing a strong correlation with laying date. Similarly, the diving effort of females was greater with increasing chick age within the same breeding stage. These results indicate that for female little penguin, the relationship between changes in prey availability and hunting effort can change at a fine scale, within a breeding stage. Therefore, any analysis of little penguin foraging behaviour during breeding should consider the timing in relation to the breeding season. We encourage researchers to develop the use of this PCA approach as it could help clarify the complexity of the underlying mechanisms determining foraging activity and we propose that it should be used as a first step of foraging behaviour analysis, before examining a particular relationship.     

An evidence of salinity impact on the bioaccumulation of heavy metals in fishes collected from the Messolonghi lagoon in Greece

The impact of salinity and water exchange rate on the accumulation of heavy metals in the benthic fish Zosterissesor ophiocephalus, collected from the shallow sublagoons Tholi and Paleopotamos, was studied. Both sublagoons are influenced by the intensive agricultural activity of the surrounding area in terms of the chemical substances’ drainage, and by the untreated sewage discharges of an urbanized area with Paleopotamos being affected in lesser extent. Paleopotamos is characterised by restricted water renovation, resulting to much higher salinity than Tholi. The bioaccumulation of the heavy metals studied (Manganese, Zinc and Copper), reveals differences among the metals with females showing higher values. Specimens from Paleopotamos showed statistically significant higher bioaccumulation of the heavy metals in both sexes, which is attributed to the extremely high salinity of this sublagoon.     

Competition among penguins and cetaceans reveals trophic cascades in the western Ross Sea, Antarctica

An apparent trophic cascade that appears during summer in the western Ross Sea, Antarctica, explains why the Antarctic silverfish (Pleuragramma antarcticum) there becomes cannibalistic; its principal prey, crystal krill (Euphausia crystallorophias) becomes scarce; and the diatom community is minimally grazed compared to adjacent areas. The krill is the major grazer of diatoms. On the basis of fieldwork at Ross Island, we suggest that the cascade results from foraging by unusually numerous Adélie Penguins (Pygoscelis adeliae), minke whales (Balaenoptera bonaerensis), and fish-eating killer whales (Orcinus orca). These species and other top predators apparently deplete the krill and silverfish. In drawing our conclusions, we were aided by two "natural experiments." In one "experiment," large, grounded icebergs altered the seasonal pattern of change in regional sea-ice cover, but not the seasonal change in penguin diet and foraging behavior that was also detected during the pre-iceberg era. In the other "experiment," a short-term polynya (opening in the ice) brought penguins and whales together in a confined area, this time altering both penguin diet and foraging behavior. We conclude that the foraging of penguins and whales, and not a formerly hypothesized seasonal decrease in sea-ice cover, explains (1) the annual switch in the penguins' prey from krill to silverfish, (2) the subsequent lengthening of penguin foraging trips, and (3) a marked decline of cetaceans in the area later in the season. Reduction in the middle-trophic-level prey is expressed in the relaxed grazing pressure on phytoplankton.     

Fatty acid signatures of stomach contents reflect inter- and intra-annual changes in diet of a small pelagic seabird,the Thin-billed prion <Emphasis Type=Italic>Pachyptila belcheri</Emphasis>

In diet analyses of seabirds, fatty acid signatures (FAS) can be used to overcome biases due to differential digestion of prey and enable the analysis of very digested diet samples. We applied FAS analysis to stomach contents of a small sub-Antarctic seabird, the Thin-billed Prion Pachyptila belcheri, which feeds mainly on squid during incubation and on crustacea during chick rearing. This seasonal dietary switch of Thin-billed prions was reflected in differences in FAS in regurgitates, as were inter-annual differences in diet composition. A discriminant function analysis correctly classified 93.4% of cases with respect to year (2006–2008) and stage of the breeding cycle (incubation versus chick rearing). The dominant types of crustacea in the diet of Thin-billed prions (amphipods Themisto gaudichaudii, euphausiids, decapods Munida gregaria, and calanoid copepods) were distinguished well by characteristic FAS patterns. However, the FAS of the two main types of prey by volume, amphipods T. gaudichaudii and squid Gonatus antarcticus, were similar to each other. Although FAS were successfully applied in the analysis of prey in stomach contents of prions, FAS of some prey species were similar and may not be distinguishable from each other if used in quantitative diet analyses.     

On the sociobiology of some hermaphroditic serranid fishes,the hamlets,in Puerto Rico

Three species of hamlets, Hypoplectrus chlorurus, H. unicolor, and H. puella (Pisces: Serranidae) were observed underwater off the west coast of Puerto Rico. They are predaceous fishes, feeding on small benthic organisms, and live in close association with the coral reef. Hamlets are highly aggressive and defend large territories, excluding their own and all other species of hamlets. They are synchronous hermaphrodites. In spawning, which occurs around sunset, one member of the pair ascends, the other joins it, each clasps the nape of the other with its tail, and gametes are released. Little color change occurs, but the black marking before the eye of H. unicolor fades when fixing on prey, proving it is not used as a sighting line. The all-black nominal species H. nigricans spawns with the yellow-tailed H. chlorurus, and is regarded here as a synonym of H. chlorurus.