Fasciole pathways in spatangoid echinoids: a new source of phylogenetically informative characters

Fascioles are important early-forming structures that play a key role in allowing irregular echinoids to burrow. They have traditionally been grouped into a small number of types according to their general position on the test, but this masks some significant differences that exist. The precise course that fasciole bands follow over the test plating has been mapped in detail for 89 species of spatangoid echinoids, representing the great majority of fasciole-bearing genera both living and fossil. Within each fasciole type, discrete and conserved patterns can be distinguished, differing both in which plates they are initiated on, and on whether they cross plate growth centres or are late-stage bands positioned towards the edge of the plate. Fasciole position is most highly conserved in the anterior and lateral interambulacral plates and on the earliest forming bands. The existence of different subanal fasciole patterns in the Micrasteridae and Brissidae suggests that these may have evolved independently. Schizasterid and hemiasterine spatangoids can each be subdivided into two major clades, and brissid spatangoids into three clades based on detailed patterns of their fascioles. Plotting fasciole pathways over test architecture provides a rich new source of phylogenetically informative characters.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 144 , 15−35.     

Tuberculation in spatangoid fascioles: Delineating plausible homologies

The existing definition of spatangoid fascioles as a narrow band of minute tubercles (miliaries) is inadequate. One reason is that many intermediates between the presence of indisputable fascioles and their total absence are known. Although fascioles are widely used in the classification of spatangoids, diversity of fasciole tuberculation has largely been ignored. We examine fascioles in about 100 spatangoid species, focusing on the earliest manifestations of fascioles (both developmentally and phylogenetically) and on their variable tuberculation. Qualitative observations are complemented by quantitative analyses (ANOVA and PCA) of 21 species. Three types of fascioles are defined according to tubercle pattern: protofascioles for localized concentrations of miliaries; parafascioles for bands of miliaries with a progressive transition to the surrounding tuberculation; orthofascioles for strongly differentiated, well-circumscribed bands of miliaries. Ontogenetic, architectural, ecological, and taxonomic variations of these three patterns are qualitatively and quantitatively investigated. We attempt to delineate plausible homologies, origins, and subsequent evolution of the different kinds of fascioles.     

Features of Transition from Larva to Juvenile in Fishes with Different Types of Early Ontogeny

The transition from the larva state to the juvenile state (i.e. morphological condition characterised by mainly adult characters) was examined in three marine fish species: herring Clupea pallasi marisalbi, wolffish Anarhichas lupus, and eelpout Zoarces viviparus, based on external morphology and skeletal development. In spite of the different reproductive styles (oviparity, facultative viviparity, and obligate viviparity, respectively) and different types of early ontogeny (indirect, transitory, and direct, respectively), the beginning of the juvenile state occurred at similar total lengths (TL), which were approximated as 35 mm TL in herring and eelpout, and 32 mm TL in wolffish. Features of ontogeny were compared, assuming that the beginning of the juvenile state represented an uniform characteristic of morphological development for these species. It was proposed that the beginning of the larva or juvenile periods (sensu Balon) could not coincide with the beginning of the larva and juvenile states in the ontogeny of some species.     

Ontogenetic patterns of relative growth in young roach Rutilus rutilus: within-river basin comparisons

To address the lack of information on the ecomorphology of young of the year juveniles (0+) freshwater fishes despite its importance to understanding geographical variation in life history, we compared the morphology of 0+ juvenile roach Rutilus rutilus from eight sub-regions of the River Great Ouse basin (U K) We tested for variation due to water retention structures, which could act to isolate upstream stocks from the general population Differences in mensural characters between the eight different regions appeared in ventral fin length and eye diameter only, which both were strongly affected by variable relative growth during early juvenile development, thus the discrepancies between regions were not due to geographical location Morphological development in roach ≥ 15 mm standard length (SL) consists of two different intervals during the first (∼ 15-40 mm SL). in contrast to adults, several mensural characters continue to develop allometrically. either positively or negatively suggesting a continuance of the larval period Whereas, m the second interval (> ∼40 mm SL). all the mensural characters except head length and eye diameter grew close to lsometrically with body length, and the rate of variation in relative growth became as strongly limited as that previously reported for adults In contrast to previous studies, which are based on few specimens we suggest this second interval to be the true start of the juvenile period     

Pourtalesiid sea urchins (Echinodermata: Echinoidea) of the northern Mid-Atlantic Ridge

Five species of deep-sea pourtalesiid echinoids were collected by the RV G.O. Sars MAR-ECO expedition to the northern part of the Mid-Atlantic Ridge: Solenocystis imitans new genus, new species, Echinosigra phiale (Thomson, 1872), Echinosigra (Echinogutta) fabrefacta Mironov, 1974, and two unidentified species of Pourtalesia. Three types of plastron plating are distinguished within the family Pourtalesiidae; in Solenocystis new genus the plating is similar to that in Spatagocystis, a genus known only from the deep-sea Antarctic. Each of three groups with distinctive plastron plating shows a continuous morphological gradation from a less elongated test without subanal rostrum to an extraordinarily elongated test with large rostrum and wide subanal fasciole; the ratio width/length of test varies from 0.86 to 0.12. Some morphological characters of the pourtalesiids are regarded as adaptations to burrowing and appear to be correlated with test elongation. The family is represented in the northern Atlantic only by 'advanced' genera (Echinosigra, Pourtalesia and Solenocystis) with very elongated tests.     

Extinction process and patterns of Middle Permian Fusulinaceans in southwest China

Study of the extinction process of 179 fusulinacean species belonging to 34 genera in southwest China demonstrates that diversity changes in the small (shell length ≤6  mm) and large (shell length >6  mm) species groups were very similar throughout the 'Maokouan' (∼Guadalupian). However, significant differences in the timing of extinction pulses occur between different fusulinacean clades with different wall structures, i.e. the nankinellids, schwagerinids, verbeekinids and neoschwagerinds, and between the large and small species groups within the schwagerinids and verbeekinids. Fusulinacean diversity reveals that the Guadalupian mass extinction began in the Middle 'Maokouan' and greatly intensified in the late 'Maokouan'. With only seven species of five genera surviving into the Late Permian, the extinction of fusulinaceans in the Guadalupian mass extinction is 96% at species level and 85% at generic level in southwest China. The preferential extinction of large, morphologically complicated species and the survival of simpler, ecologically more tolerant species of the nankinellids suggest that the extinction of Guadalupian fusulinaceans was caused by falling sea level and the consequent effects related to salinity, temperature and substrate changes. It is also demonstrated that biological characters of fusulinaceans such as shell size and test structures could have certain effects on survivorship of species in the early stages of mass extinction when extinction pressure was less intensive but were ineffectual during the extinction climax.     

The feeding ecology of juvenile mullet (Mugilidae) in south-east African estuaries

Changes in the feeding ecology of juveniles (10–59 mm standard length) of eleven species of Mugilidae were investigated in south-east African estuaries. Adults of all species spawn in the sea and juveniles enter estuaries at a length of about 10 mm. In estuaries they change their feeding habits in the following sequence: zooplankton to zooplankton in the benthos (10–15 mm), zooplankton in the benthos to meiobenthos (10–20 mm), and meiobenthos to sand particles and associated microbenthos (15–25 mm). All species show a similar pattern of change but interspecific competition is probably prevented by the rapid switch to the adult diet of microbenthos and sand particles and because species enter estuaries at different times according to spawning periods. The juveniles seek shallow quiet water areas of estuaries and it is suggested that the change from being planktonic macrophagous carnivores to benthic microphagous omnivores can only take place in the estuaries of the region. Conditions in the inshore waters of the Indian Ocean in south-east Africa are probably too rough for substrate feeding by juvenile mullet.     

Yolky eggs prepare for metabolic acceleration

Embryos convert some of their reserve into structure during development. At birth, enough must be left for further maturation, which implicitly defines the minimum amount of initial reserve. The maximum amount occurs if the mother is well-fed. Yolkiness is defined as the ratio of the maximum and the minimum amounts of initial reserve of an animal egg. Embryo development is frequently slower than can be expected on the basis of late juvenile and adult development. So development accelerates during the early juvenile stage, quantified as the ratio of the lengths at metamorphosis and at birth; during acceleration maximum specific assimilation and energy conductance increase with length. Parameters of the standard DEB model have now been estimated for some 165 species, and the goodness of fit with data is typically very good. These parameters have been used to quantify yolkiness and metabolic acceleration and a clear proportionality relationship resulted. I present some suggestions for explanations in the context of life histories.     

From cradle to early grave: juvenile mortality in European shags Phalacrocorax aristotelis results from inadequate development of foraging proficiency

In most long-lived animal species, juveniles survive less well than adults. A potential mechanism is inferior foraging skills but longitudinal studies that follow the development of juvenile foraging are needed to test this. We used miniaturized activity loggers to record daily foraging times of juvenile and adult European shags Phalacrocorax aristotelis from fledging to the following spring. Juveniles became independent from their parents 40 days post-fledging. They compensated for poor foraging proficiency by foraging for approximately 3 h d(-1) longer than adults until constrained by day length in early November. Thereafter, juvenile foraging time tracked shortening day length up to the winter solstice, when foraging time of the two age classes converged and continued to track day length until early February. Few individuals died until midwinter and mortality peaked in January-February, with juvenile mortality (including some of the study birds) five times that of adults. In their last two weeks of life, juveniles showed a marked decline in foraging time consistent with individuals becoming moribund. Our results provide compelling evidence that juveniles compensate for poor foraging proficiency by increasing foraging time, a strategy that is limited by day length resulting in high winter mortality.     

Starving early juvenile sprat Sprattus sprattus (L.) in western Baltic coastal waters: evidence from combined field and laboratory observations in August and September 2003

Growth patterns inferred from otolith microstructure analysis were compared between sprat Sprattus sprattus early juveniles (26–42 mm total length, L _T) collected in August 2003 in shallow coastal waters of the Kiel Fjord, and sprat recruits (60–95 mm L _T) sampled in October during a pelagic trawl survey of the western Baltic Sea. At the end of August, a sudden and very rapid decline in otolith growth was observed in early juveniles but not in sprat recruits. Laboratory results indicated that the early juvenile fish were starving prior to capture. Specifically, when transferred to the laboratory, otolith growth rates immediately increased in fish provided ad libitum food rations, while otolith growth of starved fish continued to decline in the same manner observed prior to field collection. In addition, the vast majority of juvenile sprat had empty stomachs on the sampling day. Given that juveniles and recruits probably experienced similar temperature conditions, the rapid decline in juvenile growth rates presumably resulted from very poor feeding conditions in nearshore waters. Starvation during the early juvenile period has not been documented before, but may, at least in the case of Baltic sprat, comprise a density-dependent mechanism operating in coastal nursery areas in some years.     

Additional observations of endobenthic behaviour in the early life history of Melanostigma atlanticum (Zoarcidae) in muddy bottom sediments of the Laurentian Trough,eastern Canada

Synopsis While adult Melanostigma atlanticum continue to be sporadically noted living within bottom sediments of the Laurentian Trough, the observation of late embryo/early juvenile stages has not previously been reported. The occurrence of 22 live specimens between 20–25 mm total length in the subsurface anoxic sediment in early May suggests that development of eggs and embryos may take place in such environments during the winter.     

Why renew fresh feathers? Advantages and conditions for the evolution of complete post‐juvenile moult

Juveniles of several passerine species renew all of their fresh juvenile feathers immediately after fledging (complete post‐juvenile moult), in contrast to the majority, which perform a partial post‐juvenile moult. To understand the adaptive roles of this phenomenon we compared the quality of juvenile plumage in species that perform a complete post‐juvenile moult with that of species which perform a partial post‐juvenile moult; we similarly compared juveniles and adults in each of these groups. The quality of feathers was measured by mass of primaries, colour, and length. In species which perform a complete post‐juvenile moult the plumage quality of second‐year individuals, in their first breeding season, is similar to the plumage quality of adults, unlike those species that perform a partial post‐juvenile moult. In species which perform complete post‐juvenile moult, the quality of the feathers grown in the nest is lower than the quality of adult post‐breeding feathers. In contrast, in species which perform partial post‐juvenile moult the quality of the feathers grown in the nest is similar to that of adult post‐breeding feathers. We found that a complete post‐juvenile moult strategy is much more common 1) in residents and short‐distance migrants than in long‐distance migrants, 2) in southern latitudes, 3) in species with medium body mass and 4) in omnivores and granivores. Our results indicate two adaptive roles of the complete post‐juvenile moult strategy: 1) achieving high quality plumage in the first year which may increase individual survival probability and fitness and 2) allocating fewer resources to nestling plumage and more to nestling development, which enables the nestlings to leave the nest earlier, thus reducing the probability of encountering nest predators. We suggest that the complete post‐juvenile moult, immediately after fledging, is an optimal strategy in favourable habitats and under low time constraints, as in some tropical ecosystems.     

External Morphology of Lophiosilurus alexandri Steindachner, 1876 during Early Stages of Development,and Its Implications for the Evolution of Pseudopimelodidae (Siluriformes)

Pseudopimelodidae are Neotropical catfishes characterized by having slightly to strongly depressed body in fully developed specimens. The largest species of the family with 500 mm SL, Lophiosilurus alexandri, experiences impressive changes in body shape during development, becoming extremely depressed when fully developed. Accordingly, Lophiosilurus alexandri is an ideal species to observe the morphological changes during ontogeny, and to seek solid interpretations on the polarity of characters. Specimens of distinct larval periods (yolk sac, flexion and postflexion; n = 186 specimens) and juvenile stages (n = 20) were analyzed. Changes in body shape, position of mouth and eye, morphology of fins and pigmentation were observed during the development of Lophiosilurus. Larvae (5.7–11.2 mm standard length) had pigmentation concentrated on the head and parts of body, eyes small and pigmented, short barbels, and well-developed finfold. Juveniles (15.9–28.1 mm standard length) had body shape similar to adult, with head depressed and bearing bony ridges, large mouth, dorsally-oriented eyes, small barbels and well-developed shoulder bulges (cleithral width). The greatest morphological changes in the development of L. alexandri occurred during the postflexion larval stage. Relative to standard length, measurements of snout length, head depth and body depth are smaller in juveniles than in larvae, but body width is larger. New interpretations on the phylogenetic characters related to these changes are provided in view of the two alternative hypotheses of the evolution of Pseudopimelodidae.     

Fledgling survival increases with development time and adult survival across north and south temperate zones

Slow life histories are characterized by high adult survival and few offspring, which are thought to allow increased investment per offspring to increase juvenile survival. Consistent with this pattern, south temperate zone birds are commonly longer‐lived and have fewer young than north temperate zone species. However, comparative analyses of juvenile survival, including during the first few weeks of the post‐fledging period when most juvenile mortality occurs, are largely lacking. We combined our measurements of fledgling survival for eight passerines in South Africa with estimates from published studies of 57 north and south temperate zone songbird species to test three predictions: (1) fledgling survival increases with length of development time in the nest; (2) fledgling survival increases with adult survival and reduced brood size controlled for development time; and (3) south temperate zone species, with their higher adult survival and smaller brood sizes, exhibit higher fledgling survival than north temperate zone species controlled for development time. We found that fledgling survival was higher among south temperate zone species and generally increased with development time and adult survival within and between latitudinal regions. Clutch size did not explain additional variation, but was confounded with adult survival. Given the importance of age‐specific mortality to life history evolution, understanding the causes of these geographical patterns of mortality is important.     

Intra-plant host selection, oviposition preference and larval survival of Chrysophtharta agricola (Chapuis) (Coleoptera: Chrysomelidae: Paropsini) between foliage types of a heterophyllous host

Abstract 1 Paropsine chrysomelid beetles defoliate commercial eucalypt plantations in Australia. Adults and larvae feed on the same host, with the larval food source determined by the oviposition choice of females. Most eucalypt species are heterophyllous, with their foliage undergoing distinct morphological and chemical changes between adult and juvenile growth. 2 The intra‐plant foliage feeding and oviposition preference adults and the larval development of Chrysophtharta agricola were examined using adult and juvenile foliage of a heterophyllous plantation species, Eucalyptus nitens. The foliage types differ in chemistry, toughness, waxiness and timing of production. 3 In the field, feeding damage caused by adult beetles was 15% more frequent on adult foliage than on juvenile foliage; however, egg batches were three times more common on juvenile than on adult foliage. 4 Oviposition preference for juvenile foliage over adult foliage was confirmed in choice trials in the laboratory, with adult fecundity and longevity not significantly different between foliage types. 5 Larval survival, development time and subsequent pupal weight were also unaffected by foliage type, suggesting that neither foliage type is nutritionally superior for adults or for larvae. However, adult foliage was significantly thicker than juvenile foliage and this may prove a physical constraint to larval establishment. Biotic and abiotic factors (including interactions with natural enemies, competition, microclimate and mate location) that may affect patterns of host plant utilization are discussed.     

Benthic ctenophore (Order Platyctenida) reproduction,recruitment, and seasonality in south Florida

Reproductive structures, modes, and seasonal patterns of size–class abundances are examined in two benthic platyctene (Family Coeloplanidae) ctenophore species present in dissimilar shallow marine environments in subtropical southeast Florida. Coeloplana waltoni, a minute (1–3 mm body length) epizoic associate of octocorals, occurs in exposed environments often under turbulent conditions, and Vallicula multiformis (2–10 mm) commonly occurs epiphytically on macroalgae in protected, calm‐water environments. Reproductive activity in C. waltoni is most active during the warm‐water summer season (June–October); gonadal development in V. multiformis occurs year‐round, and is most pronounced during sea‐warming periods in late spring (May) and late summer to early autumn (August–October), with release of cydippid larvae. Both species are hermaphroditic brooders, exhibiting paedogenesis (early gonadal development) at body lengths approximately one‐third (Coeloplana) to one‐sixth (Vallicula) of maximum adult size. Juvenile individuals (<0.6 mm) increased in abundance in C. waltoni during the summer reproductive period, and large (≥1 mm) pink‐colored individuals comprised 50% or more of samples from July through September. Seasonal abundance of gravid individuals and the timing of cydippid larval release in V. multiformis did not correspond closely with juvenile or adult population densities. Asexual fragmentation occurred in both ctenophore species, but was observed more frequently in individuals of V. multiformis. This asexual mode of reproduction probably accounted in part for the discordance between ctenophore abundances and larval recruitment events by sexual means. Morphological structures and behaviors associated with reproduction are described in this study. Uncommon images of reproductive products (gametes, embryos, larvae), spawning events, brooding, and asexual fragmentation are included, some for the first time in the published literature.     

Shoaling in juvenile guppies: the effects of body size and shoal size

While factors affecting shoal mate choice have been examined extensively in adult guppies (Poecilia reticulata), few studies have focused on the shoaling behavior of juveniles. In this study, juvenile guppies were tested for their ability to shoal as well as their response to shoal mates of different body size and to shoals with different numbers of individuals. In dichotomous choice tests, 10-day-old guppies (mean body length=8.83 mm), 30-day-old guppies (13.17 mm) and 50-day-old guppies (18.6mm) were given the opportunity to swim near shoals of five fish or an empty chamber. In most cases, the juvenile fish demonstrated shoaling behavior, swimming near a group of fish rather than an empty chamber, regardless of the age of the stimulus shoal. When presented with two shoals, one of similar age and body size and one of dissimilar age and body size, only the 50-day-old guppies showed a significant preference for the age-matched shoal. Similarly, when choosing between a large shoal and a small shoal, only the 50-day-old guppies spent significantly more time near the larger shoal. Thus, while juveniles at each age shoaled, only 50-day-old fish demonstrated the shoal mate discrimination seen in adult fish.     

Spatial Variation in Abundance,Size and Orientation of Juvenile Corals Related to the Biomass of Parrotfishes on the Great Barrier Reef,Australia

For species with complex life histories such as scleractinian corals, processes occurring early in life can greatly influence the number of individuals entering the adult population. A plethora of studies have examined settlement patterns of coral larvae, mostly on artificial substrata, and the composition of adult corals across multiple spatial and temporal scales. However, relatively few studies have examined the spatial distribution of small (≤50 mm diameter) sexually immature corals on natural reef substrata. We, therefore, quantified the variation in the abundance, composition and size of juvenile corals (≤50 mm diameter) among 27 sites, nine reefs, and three latitudes spanning over 1000 km on Australia’s Great Barrier Reef. Overall, 2801 juveniles were recorded with a mean density of 6.9 (±0.3 SE) ind.m⁻², with Acropora, Pocillopora, and Porites accounting for 84.1% of all juvenile corals surveyed. Size-class structure, orientation on the substrate and taxonomic composition of juvenile corals varied significantly among latitudinal sectors. The abundance of juvenile corals varied both within (6–13 ind.m⁻²) and among reefs (2.8–11.1 ind.m⁻²) but was fairly similar among latitudes (6.1–8.2 ind.m⁻²), despite marked latitudinal variation in larval supply and settlement rates previously found at this scale. Furthermore, the density of juvenile corals was negatively correlated with the biomass of scraping and excavating parrotfishes across all sites, revealing a potentially important role of parrotfishes in determining distribution patterns of juvenile corals on the Great Barrier Reef. While numerous studies have advocated the importance of parrotfishes for clearing space on the substrate to facilitate coral settlement, our results suggest that at high biomass they may have a detrimental effect on juvenile coral assemblages. There is, however, a clear need to directly quantify rates of mortality and growth of juvenile corals to understand the relative importance of these mechanisms in shaping juvenile, and consequently adult, coral assemblages.     

Neuregulin‐1 is essential for nerve plexus formation during cardiac maturation

The Neuregulin‐1 (Nrg1)/ErbB pathway plays multiple, critical roles in early cardiac and nervous system development and has been implicated in both heart and nerve repair processes. However, the early embryonic lethality of mouse Nrg1 mutants precludes an analysis of Nrg1's function in later cardiac development and homeostasis. In this study, we generated a novel nrg1 null allele targeting all known isoforms of nrg1 in zebrafish and examined cardiac structural and functional parameters throughout development. We found that zebrafish nrg1 mutants instead survived until young adult stages when they exhibited reduced survivorship. This coincided with structural and functional defects in the developing juvenile and young adult hearts, as demonstrated by reduced intracardiac myocardial density, cardiomyocyte cell number, swimming performance and dysregulated heartbeat. Interestingly, nrg1 mutant hearts were missing long axons on the ventricle surface by standard length (SL) 5 mm, which preceded juvenile and adult cardiac defects. Given that the autonomic nervous system normally exerts fine control of cardiac output through this nerve plexus, these data suggest that Nrg1 may play a critical role in establishing the cardiac nerve plexus such that inadequate innervation leads to deficits in cardiac maturation, function and survival.     

The growth pattern of chimpanzees: Somatic growth and reproductive maturation inPan troglodytes

Growth of chimpanzees reared at the Kumamoto Primates Park of Sanwa Kagaku Kenkyusho Co. Ltd. was studied cross-sectionally from the viewpoints of somatic growth and reproductive maturation. Distance and velocity curves were expressed using spline function method. Males showed adolescent growth acceleration in body weight, with a peak at 7.86 yrs of age, but not in trunk length. Females showed continuous rapid growth from mid-juvenile to adolescent phase in both body weight and trunk length, but no isolated adolescent spurt. The Sanwa chimpanzees matured at about 12.5 yrs of age for females and 15.0 yrs for males. The mean adult weights and trunk lengths were 53.2 kg and 507.8 mm for males and 42.7 kg and 481.6 mm for females. The Sanwa chimpanzees had similar growth patterns to those of the Yerkes chimpanzees, although they showed a slight delay in infancy, and a higher growth rate from the early juvenile phase onwards. Growth patterns in these two laboratories may be regarded as “normative” for laboratory-reared chimpanzees. They matured earlier than wild chimpanzees by more than two years. The major reason for the retarded maturation in wild chimpanzees is the delay of growth from infant to the early juvenile phases (0–4 yrs of age), probably owing to a limited nutritional supply from the mother. Development of the testes comprised three phases: slow growth from infant to juvenile (until 6.4 yrs); rapid growth around adolescence (until 9.2 yrs); and adult (mean testicular volume, 187 cm³). Setting the nutritional standard at 2,000–2,600 Cal/day (= Kcal/day) per adult, calories were considered for captive chimpanzees in each age class.