Seasonal cambium activity in the subtropical rain forest tree Araucaria angustifolia

Information on the timing and dynamics of tree ring formation is essential to assess the seasonal behavior of secondary wood growth and its associated environmental influences. Araucaria angustifolia is a dominant species in highland pluvial ecosystems of southeastern South America. Previous investigations indicated that their growth rings are formed annually, but no information exists about the timing of growth ring formation and the environmental triggers influencing cambium activity. In this paper we examine inter- and intra-annual cambial activity in A. angustifolia, through anatomical and dendrochronological evidence at two study sites, and model the relationships between regional climate variation and intra-annual tree ring formation. The results confirm the annual growth ring formation in A. angustifolia and indicate that its growth season extends from October to April. Day length and temperature were the main environmental factors influencing the seasonal cambium activity. Our results evidence the dendrochronological potential of A. angustifolia for ecological and climatological studies in southeastern South America.     

Mangrove growth rings: fact or fiction?

The analysis of tree rings in the tropics is less straightforward than in temperate areas with a demarcated unfavourable winter season. But especially in mangroves, the highly dynamic intertidal environment and the overriding ecological drivers therein have been a reason for questioning the existence of growth rings. This study aimed at casting light on growth rings in mangroves. In six mangrove species growing in Gazi Bay, Kenya (Sonneratia alba, Heritiera littoralis, Ceriops tagal, Bruguiera gymnorrhiza, Xylocarpus granatum and Lumnitzera racemosa), the occurrence of growth rings was examined. Growth rate of each species was determined based on a 1-year period using the cambial marking technique. The effect of climate was furthermore considered by comparing the results with a number of wood samples originating from contrasting climatic regions. We can conclude that for growth rings to appear in mangroves more than one condition has to be fulfilled, making general statements impossible and explaining the prevalent uncertainty. Climatic conditions that result in a range of soil water salinity experienced over the year are a prerequisite for the formation of growth rings. For species with an anatomy characterized by indistinct ring boundaries, this should be combined with a growth rate of at least 0.3 mm/year. The use of growth rings for age or growth rate determinations should thus be evaluated on a case by case basis.     

Annual Growth Ring Patterns in Brachystegia spiciformis Reveal Influence of Precipitation on Tree Growth1

The availability of exactly dated tree‐ring chronologies is limited in tropical regions. However, these chronologies could contribute widely to studies of the influence of natural and human‐induced factors on tropical forests. We examine the potential for building a chronology based on three sites in the miombo woodland of western Zambia. Brachystegia spiciformis Benth., a dominant species from this vegetation type, is used. Response of the chronology to several climatic factors is examined. All specimens showed very clear growth rings, and cross‐dating between radii of a tree was successful for all trees. Site chronologies could be constructed after cross‐dating of growth ring series of individual trees. The mean growth ring curves of the three sites were significantly similar, allowing for the construction of a regional chronology. Correlation function analysis between the tree‐ring chronology and regional climatic variables revealed that climate at the core of the rainy season, in December and January, has an explicit influence on tree growth. Where precipitation and relative humidity in these months influence tree growth positively, temperature correlates in a negative way. Some 20 percent of the variance in the B. spiciformis tree‐ring chronology is accounted for by wet season rainfall. The successful cross‐dating and correlation between a tree‐ring chronology and climate demonstrated in this study indicate annual ring formation in B. spiciformis trees and sensitivity to climatic conditions.     

On the reliability of the growth rings in two species of the family Characidae in the Sudan

The scales, opercula and vertebrae from 69 Hydrocyon forskali and 63 Alestes dentex were investigated for their reliability as aging structures. In both species the scales were unreliable due to their regenerative property, in spite of the clarity of their growth rings.Back-calculated lengths from the first growth ring laid on opercula and vertebrae resulted in values that coincided with the annual growth of the two species. In H. forskali, the mean back-calculated length from the two bones were 16.2 ± 5.9 and 16.8 ± 4.4 respectively, and for A. dentex, 11.1 ± 1.7 and 12.8 ± 1.8, respectively.Predictive equations for bone radius from fish length were written for both species. In this study, reliability was maximal between the observed and back-calculated lengths for both species (r > 0.95).     

SPACING OF PARENCHYMA BANDS IN WOOD OF CARYA GLABRA (MILL.) SWEET,PIGNUT HICKORY,AS AN INDICATOR OF GROWTH RATE AND CLIMATIC FACTORS

Variations in the spacing of concentric, tangential bands of apotracheal axial wood parenchyma cells in the annual growth rings of Carya glabra (Mill.) Sweet constitute a record of changes in radial growth rate and of fluctuations in climatic factors. This relationship was determined from trees that were growing under moisture stress and consequently had highly consistent ring-width patterns from tree to tree. Band spacing decreased across a growth ring and was generally greater in wide than in narrow rings. Thus, the proportion of xylem cross-sectional area occupied by axial parenchyma tissue was inversely related to ring width. Average numbers of bands in each of 67 years were analyzed relative to ring widths. Years in which bands were unusually numerous relative to ring width were years of low April-May precipitation, leading to slow spring growth and closely spaced bands early in the growing season. Years in which band numbers were unusually low relative to ring width were years of high April-May precipitation, leading to rapid growth and widely spaced bands early in the season; low July precipitation in these years apparently curtailed growth before much of the typical late-season wood, with its closely spaced bands, could form. Parenchyma bands were aggregated within some rings, suggesting that a slowing of growth had been followed by a growth spurt. Evidence for a late-season spurt was that rings with aggregations were significantly wider than rings formed the same year and not having aggregations of bands. The number of vessels per unit of cross-sectional area was inversely related to ring width.     

Common otolith microstructure related to key early life‐history events in flatfishes identified in the larvae and juveniles of cresthead flounder Pseudopleuronectes schrenki

Otolith microstructure of reared and wild cresthead flounder Pseudopleuronectes schrenki larvae and juveniles was used to investigate the daily periodicity of ring formation, morphological change and unique otolith structure related to important life events. By comparing microstructural features of P. schrenki with those reported for other flatfish species, it was shown that there may be microstructural features that are common to all flatfishes. In the sagittae and lapillus, a check (a distinct ring) was formed in the centre of otoliths at c. 6 days post hatching, and the daily formation of rings observed outside the check was confirmed. During metamorphosis, accessory primordia (AP) of otolith growth were formed on the outer edge of the sagittae, and the shape of the sagittae became more complex. No AP was formed on the lapilli, however, and otolith rings were concentrically formed throughout the larval and juvenile (≤51·6 mm standard length, L_S) stages. It is proposed, therefore, that lapilli are more appropriate than sagittae for analysis throughout the larval and juvenile (≤51·6 mm L_S) stages. During metamorphosis, unique rings that are relatively wide and show weak contrast are formed on lapilli (metamorphosing zone, MZ). Hence, the duration of metamorphosis, larval duration and the days of juvenile life can be estimated by the number of rings within the MZ, using rings from the check to outermost ring of the MZ, and that of rings formed outside MZ, respectively. The formation of AP on sagittae as well as the absence of AP, bilateral asymmetry and the formation of a unique structure during metamorphosis on lapilli have also been reported for other flatfishes.     

The record of daily growth in otoliths of Atlantic silversides, Menidia menidia, from field and laboratory

Otoliths were removed from field-collected silversides of age less than 3 months. Otolith diameter was highly correlated with total length of the fish. Daily growth ring counts for this species are known to be a function of age rather than size, so widths for the daily growth rings provide a record of daily increases in length of the fish. Measurement of ring widths showed that weekly specific growth rate was greater than 70% at age 1 week, but declined to about 30% at age 1 month and about 15% at age 2 months. A laboratory experiment in which temperature was changed on a weekly basis demonstrated that environmental variables can affect the width of rings. Nevertheless, the growth rate of field-collected fish, as calculated from otolith ring widths, was more highly correlated with size of fish, as measured by otolith radius, than with the environmental variables of temperature, salinity and plankton abundance. Back-calculation of growth rates from otolith ring widths of five fish collected at the end of the growing season yielded the same age-growth curves as were obtained from 203 fish collected biweekly during the season.     

Growth-ring variations of dwarf shrubs reflect regional climate signals in alpine environments rather than topoclimatic differences

Aim Our main aim is to determine if ring‐width variations in Empetrum hermaphroditum reflect regional or local topoclimate signals in an alpine environment. In the case that topoclimate provides the dominant signal, a secondary aim is to link these to spatial distribution patterns of different vegetation types. Location The study area is situated in the middle alpine belt in the Vågåmo region, Central Norwegian Scandes. Sampling sites cover different topoclimates: ridges, north‐facing slopes and south‐facing slopes. Methods We constructed ring‐width chronologies of E. hermaphroditum for each type of microsite for the common period 1951–2004. Climate data were prepared on an hourly, daily and growing‐season time scale. Climate–growth relationships were evaluated using bivariate correlations and regression tree methods for continuous time‐series analyses. In addition, extreme growth anomalies (pointer years) were compared with the climate conditions in those years. The impact of water supply on wood anatomy was determined by correlating the conductive area (percentage of vessel per growth ring) with a running mean (sum) of 10‐day intervals for temperature and precipitation. Results This study indicates that mean summer (June–August) temperatures determine the width of the growth rings of E. hermaphroditum irrespective of topoclimate. The length of the growing season, which is the most differentiating climatic factor between microsites, does not substantially alter the anatomical ring structure. Microsite differences in mean growth rates are attributed to the higher frequency of warm days. Extremely warm days limit ring‐width development at south‐facing slopes, while plants at ridges and north‐facing slopes still benefit from higher temperatures. As a consequence, pointer years are not developed synchronously at all microsites. Vessel formation is affected by available moisture, especially in the later part of the growing season. Main conclusions Topoclimate induces slight modifications of annual growth‐ring increments of E. hermaphroditum at different microsites. In contrast to the distribution patterns of vegetation types that are determined by snow cover, growth‐ring variations are related to summer temperature conditions, and the prominent regional climate signal is still reflected at all microsites. This offers the opportunity to reconstruct climatic change in alpine regions from dwarf shrub ring‐width chronologies.     

Age determination in the sea urchin paracentrotus lividus (Lamarck), with notes on the reproductive cycle

Evidence from other species of sea urchins indicated that the growth rings found in the skeletal plates of Paracentrotus lividus (Lamarck) were probably formed annually, and could therefore be used as a means of determining age. In this species, annual formation of the rings has been proved by the similarity of growth curves derived from ring counts and size-frequency analyses; there is a good agreement between the annual increments predicted from these growth curves and those observed in experimental populations. Determinations of the cycles of ring deposition and reproduction gave results which were consistent with this interpretation.Similarities in the growth curves did not always extend to the mean values determined for each ring-class and the equivalent size-frequency groups, and the size-frequency distributions of single ring-classes were found to be much wider than those of the equivalent groups determined by sizefrequency analysis. This is attributed to a wide variation in the season of ring deposition in different individuals, and this limits the accuracy of ring count ageing to about ± one year.There were no significant differences between the reproductive cycles of male and female urchins; the gonad indices rose to peak values in late winter and late summer. Gonad condition was very variable within each population sample. The animals grew to a mean diameter of 35–50 mm in about four years, after which growth rates slowed down, and it is probable that most of the urchins died of senescence after 6–9 years.     

The growth of Tilapia mossambica Peters (Pisces: Cichlidae) in Lake Sibaya, South Africa

The scales of Tilapia mossambica Peters * from Lake Sibaya show clear rings formed by broken, widely-spaced circuli in the anterior field. In order to determine the rate and time of scale ring formation, the number of circuli in the marginal increment was determined. Five scales from the pectoral region of 2223 fishes caught at regular intervals over two years were examined. When plotted as histograms, the number of circuli in the marginal increment showed a sharp movement to the left in September–October and again in January, indicating that the rings were formed during these months in the majority of fish examined. Pre- and postbreeding feeding migrations, and increases in condition factor, were associated with scale ring formation. Otoliths and opercula were also examined for age determination. Annual length increments were calculated for 450 T. mossambica collected throughout the year, and growth curves plotted. T. mossambica from Sibaya reach maturity after one year at a standard length of about 8 cm in females, and after two years at 10 cm in males. The breeding population had a standard length mode of 11–12 cm in females and 17 cm in males. The maximum final size was about 24 cm.     

Otolith growth and age estimation in the European hake

The internal sulcal rings in Merluccius merluccius otoliths cannot be considered as annuli. In the absence of a strong seasonal Zeitgeber, hake otoliths in the Mediterranean did not lay down an interpretable ring pattern that would be useful for age determination. A total of 484 sagittal otoliths from specimens ranging between 6 and 94 cm L T was studied in monthly samples from the Gulf of Lions in 1989–1990. Transverse, burnt otolith sections was analysed with an image analysis system using enhanced and filtered images and using a fast Fourier transform (FFT) function to avoid subjectivity in ring interpretation. The otolith radius-fish length relationship showed allometric growth and sexual dimorphism. The ring pattern was consistent for the sulcal rings in both sexes. Changes in the marginal increment showed the formation of multiple sulcal rings, of both environmental and physiological origin. The ring pattern depended on the sex and sexual activity.     

Seasonal dimorphism in wood anatomy of the Mediterranean Cistus incanus L. subsp. incanus

In Mediterranean-type ecosystems, seasonal dimorphism is an adaptive strategy aimed to save water by developing brachyblasts with xeromorphic summer leaves as opposed to dolichoblasts with more mesomorphic winter leaves. The aim of this study was to analyse the anatomical properties of 1-year-old twigs of Cistus incanus subsp. incanus, a seasonally dimorphic shrub, to highlight properties allowing its adaptation to the Mediterranean environment. A more specific purpose was to verify the occurrence of seasonal dimorphism in wood anatomy in order to understand: (a) whether and to what extent the traits of efficiency/safety of water transport are expressed in brachyblasts and dolichoblasts, and (b) the effects on the formation of growth ring boundaries in wood. Our overall analysis showed that anatomical features of branches in C. incanus are designed to: (a) protect from desiccation by developing thick cuticle and suberized epidermal and sub-epidermal layers; (b) defend the plant from predators by accumulating phenolics; and (c) regulate water transport through the development of specific wood anatomy, according to the season, thus optimising properties of efficiency/safety. Regarding the latter point, our results indicated that brachyblast wood is safer than dolichoblast wood insofar as it has narrower and more frequent vessels; measurement of other specific anatomical traits, such as vessel wall thickness, suggested that brachyblast wood has a higher resistance to implosion due to drought-induced embolism. Finally, peculiar anatomy of brachyblast and dolichoblast wood results in the formation of so-called false rings. Hence, wood rings in C. incanus should be considered to be “seasonal” rather than “annual”.     

Age and Long-term Growth of Trees in an Old-growth Tropical Rain Forest, Based on Analyses of Tree Rings and 14C1

In an old‐growth tropical wet forest at La Selva, Costa Rica, we combined radiocarbon (¹⁴C) dating and tree‐ring analysis to estimate the ages of large trees of canopy and emergent species spanning a broad range of wood densities and growth rates. We collected samples from the trunks of 29 fallen, dead individuals. We found that all eight sampled species formed visible growth rings, which varied considerably in distinctiveness. For five of the six species for which we combined wood anatomical studies with ¹⁴C‐dates (ring ages), the analyses demonstrated that growth rings were of annual formation. The oldest tree we found by direct ring counting was a Hymenolobium mesoamericanum Lima (Papilionaceae) specimen, with an age of ca. 530 years at the time of death. All other sampled individuals, including very large trees of slow‐growing species, had died at ages between 200 and 300 years. These results show that, even in an everwet tropical rain forest, tree growth of many species can be rhythmic, with an annual periodicity. This study thus raises the possibility of extending tree‐ring analyses throughout the tropical forest types lacking a strong dry season or annual flooding. Our findings and similar measurements from other tropical forests indicate that the maximum ages of tropical emergent trees are unlikely to be much greater than 600 years, and that these trees often die earlier from various natural causes.     

The use of X-ray densitometry to evaluate the wood density profile of Tectona grandis trees growing in fast-growth plantations

Tectona grandis (teak) is an important commercial tree species that is widely used in tropical dendrochronology due to the formation of climate-sensitive annual growth rings. However, young trees growing in plantation conditions exhibit poor ring visibility during the first years of growth, limiting the dendrochronology application. In the present study, we use x-ray densitometry to determine the wood density profile between and within annual rings and at the sapwood-heartwood boundary in trees from fast-growth plantations. The resulting wood density profiles (WDP) can be categorized as uniform, stable growth, unstable growth, and false. The annual ring boundaries were indistinct in trees less than 8 years old. In mature trees, the annual ring boundaries are more defined. In relation to the sapwood-heartwood boundary, the WDP showed a decrease in the wood density; however, this decrease is influenced by the annual ring boundary when the two boundaries coincide. The identification of annual rings in trees growing in fast-growth plantations should be combined with X-ray densitometry and visual identification if wood density data are necessary for deriving other analysis, as climate change, from annual ring.     

Relating tree growth to rainfall in Bolivian rain forests: a test for six species using tree ring analysis

Many tropical regions show one distinct dry season. Often, this seasonality induces cambial dormancy of trees, particularly if these belong to deciduous species. This will often lead to the formation of annual rings. The aim of this study was to determine whether tree species in the Bolivian Amazon region form annual rings and to study the influence of the total amount and seasonal distribution of rainfall on diameter growth. Ring widths were measured on stem discs of a total of 154 trees belonging to six rain forest species. By correlating ring width and monthly rainfall data we proved the annual character of the tree rings for four of our study species. For two other species the annual character was proved by counting rings on trees of known age and by radiocarbon dating. The results of the climate–growth analysis show a positive relationship between tree growth and rainfall in certain periods of the year, indicating that rainfall plays a major role in tree growth. Three species showed a strong relationship with rainfall at the beginning of the rainy season, while one species is most sensitive to the rainfall at the end of the previous growing season. These results clearly demonstrate that tree ring analysis can be successfully applied in the tropics and that it is a promising method for various research disciplines.     

Assessing the utility of direct and indirect methods for estimating tropical tree age in the Western Ghats, India

The knowledge of tree age is important for understanding tree growth and forest dynamics. It may be estimated by ‘direct’ methods involving growth ring counts, or by ‘indirect’ methods involving field measurements of growth rates. Direct methods are considered more accurate, but it is not clear if they are appropriate for all species, notably from the humid tropics. In this paper we assess the occurrence of annual growth rings and their utility for age estimation in three tropical tree species, Acrocarpus fraxinifolius, Dalbergia latifolia (Fabaceae) and Syzygium cumini (Myrtaceae), growing in traditional shade coffee plantations of the southern Western Ghats, India. These species previously were described as having “indistinct or absent” growth rings. We used anatomical studies, field measurements and computational methods to characterise growth rings and assess similarities between directly and indirectly estimated tree ages. Our study revealed that annual growth rings were characterised by different sets of anatomical features per species and were most distinct in the fast-growing deciduous A. fraxinifolius. Growth rates measured in the field showed annual periodicity in all three species, and reflected annual rainfall-drought cycles in D. latifolia and S. cumini. Direct age estimates were most similar to indirect estimates in D. latifolia, and least so in S. cumini. The results of direct age estimation by counting rings are consistent with them being annual in nature in tropical species with distinct and reliable annual growth ring formation. However, for species with poorly defined growth rings, indirect age estimation methods might be more useful.     

Orderly formation of the double ring structures for plastid and mitochondrial division in the unicellular red alga Cyanidioschyzon merolae

The formation of the plastid-dividing ring (PD ring) and mitochondrion-dividing ring (MD ring) was studied in a highly synchronous culture of the unicellular red alga Cyanidioschyzon merolae. The timing and the order of formation of the MD and PD rings were determined by observing organelles around the onset of their division, using transmission electron microscopy. In  C. merolae, there is one chloroplast and one mitochondrion per cell, and the shape of the chloroplast changes sequentially from acorn-like, to round, to trapezoidal, to peanut-shaped, in that order, during the early stage of chloroplast division. None of the cells with acorn-shaped or round chloroplasts contained organelles with PD rings or MD rings, while all of the cells with peanut-shaped chloroplasts contained organelles with both PD rings and MD rings. In cells with peanut-shaped chloroplasts, the PD and MD rings were double ring structures, with an outer ring located on the cytoplasmic face of the outer membrane of the organelle, and an inner ring located in the matrix beneath the inner membrane. These results suggested that the double ring structures of the PD ring and the MD ring form when chloroplasts are trapezoidal in shape. Detailed three-dimensional observation of cells with trapezoidal chloroplasts revealed the following steps in the formation of the double ring structures of the PD and MD rings: (i) the inner ring of the PD ring forms first, followed by the outer ring; (ii) then the MD ring forms and becomes visible; (iii) when the double ring structures of the two rings have formed, the microbody then moves from its remote location to the plane of division of the mitochondrion and contraction of the PD and MD rings commences. These steps were also confirmed by computer-aided three-dimensional reconstruction of the images from serial thin sections. This study reveals the order of formation of the double ring structures of the PD and MD rings, and the behavior of the microbody around the onset of division of plastids and mitochondria. The results also provide the first evidence that the inner PD ring is not a tension element formed by the contractile pressure but a definite structure, independent of the outer ring. Received: 31 March 1998 / Accepted: 14 May 1998     

Evoluzione Annuale della Cerchia Legnosa in Individui Maschili E Femminili di Ginkyo Biloba L.

Abstract

Wood ring evolution in male and female plants of GINKYO BILOBA L. — A morphological examen of the wood rings in Ginkyo biloba L. has been performed on stem sections from specimens of the Botanical Institute of Florence, on wood samples from the collection of Adriano Fiori and on wood cores extracted from the stem of male and female plants growing in the Botanical Garden of Florence.

At the same time the development of the growth ring has been followed on wood samples tacken off the stems of two adult Ginkyo plants growing in the Botanical Garden of Florence.

All the material which has been studied shows the same characters, that is the frequency of uncomplete crescent-shaped wood rings, and consequently a large degree of eccentricity of the stems, depending not only upon the uncomplete wood rings, but also on the changing width of each ring.

Microscopical analysis has pointed out the presence of false rings, limited by discontinuous late wood, particularly in the female plant. This character has been observed only in a fossil wood of Ginkyo bonesii.

It has not been possible to assers how many rings or crescents are formed during one year. From dating records made on wood cores it seems that more than one ring could differentiate during one year, but the cambium shows a very irregular activity in Ginkyo. The factors which control the formation of false and uncomplete wood rings (wood crescents) are also unknown; their formation possibly depending upon environment conditions.

External factors such as water deficiency probably control cambial activity through auxin production. Larson (1964) has shown that draught causes a decrease in tracheids diameter, and the formation of false growth rings; thus the same causes could have been in action in our plants.

Noteworthy are also the data by Gungkel and Thimann (in Larson 1964) showing that auxin content increases downwards in the shoots of Ginkyo and that the apical bud has a very low auxin production, so that it seems possible that in Ginkyo the leaves supply something which the cortical tissue converts into auxin.

Lastly, the almost continuous growth activity of the cambium in Ginkyo must be pointed out, this causes a great difficulty in checking the beginning and the end of this process along the year. It is any way possible to establish the time of the highest cambial activity, which lies in June-July both in the female and in the male plants.     

Effect of climate on the growth of annual rings in the main roots of perennial forbs in an Inner Mongolian semi‐arid grassland,China

Question: Is there a pattern in growth of annual rings in roots of perennial forbs in relation to climate and climate extremes in grassland ecosystems? Location: Semi‐arid grassland in Duolun (42°27′N, 116°41′E, 1380 m a.s.l.), central Inner Mongolia, China. Methods: Main roots of three perennial species, Potentilla anserina L., Cymbaria dahurica L. and Lespedeza daurica Schindl., were sampled. Cross‐sections (10–15‐μm thick) were produced from the proximal end of sampled roots using a sledge microtome. Annual growth rings in the main roots were identified and measured by differentiating between earlywood and latewood in the secondary xylem. Relationships between annual growth rings and monthly mean temperature and total monthly precipitation were identified using correlation analysis. Differences in an annual ring width to the previous and following years were examined by calculating a distinctness score. Results: The three perennial forbs showed clearly demarcated annual growth rings in all individuals and the same fluctuation patterns. Their ring widths were generally positively correlated with precipitation from April to October (except for August) and with temperature from February to June (except June for L. daurica), September to October, and the annual mean. Strong deviations of annual ring widths from their neighbour rings were observed in 1998 and 2000. The trend of absolute distinctness scores (D_m) increased significantly from 1988 to 2003, indicating an increase in the frequency of annual ring width variation. Conclusions: Annual growth rings in the main roots of three perennial forb species can be used as an indicator of the influence of climate on below‐ground grassland growth. The change in below‐ground conditions and effects on the functioning of grassland should receive more attention in future studies.     

Some aspects of the biology of the horse mackerel [Trachurus trachurus (L.)] in waters around Britain

Evidence is presented which supports the hypothesis of a seasonal migration between the North Sea and the western English Channel. Fish caught west of Britain have a different length composition and there is some evidence of a North–South migration. Analysis of stomach contents shows that the species is a plankton feeder. Data on seasonal growth in young fish show the growth-rate is most rapid between August and December; length-frequency distributions of two year-classes were bimodal. The use of the otolith burning technique revealed up to 35 check rings, as compared with the reported maximum age of 10 years. With the exception of ring numbers 1 to 3, it could not be established the rings were annual.