Larval growth in the river lamprey, Lampetra fluviatilis

A series of 987 ammocoetes from the rivers Towy, Teme, and Taw have been identified as mainly L. fluviatilis (L.) on the basis of oocyte counts on female ammocoetes. The length frequency distributions for this material differs from either L. planeri or P. marinus in showing only three modes in addition to the young of the year and the length distribution of the final mode coincides with the length range for 119 metamorphosing and macrophthalmia stages of L. fluviatilis that have been found at the same sites. These animals measured from 80–117 mm in length and weights varied from 0.76–2.28 g. Metamorphosis is believed to take place in late summer and early autumn when in the majority of cases, the ammocoetes are four and a half years old. The evidence that the non-parasitic L. planeri has a longer larval life than the closely related parasitic L. fluviatilis is thought to have some significance in relation to the evolution of the brook lamprey species.     

A comparison of the metamorphosing and macrophthalmia stages of the lampreys Lampetra fluviatilis and L. planeri

Comparisons of metamorphosing and macrophthalmia stages of the closely related species, L. fluviatilis (L.) and L. Planeri (Bloch), have shown that these can be distinguished within a few weeks of the onset of metamorphosis by characteristic differences in colouration and body form. Measurements of several body intervals have disclosed differences between the macrophthalmia stages of the two species. A sharp distinction between the blunt teeth of L. planeri and the supposedly sharp teeth of L. fluviatilis has not been confirmed in these early stages, but significant differences have been found in the numbers of teeth in the anterior field of the oral disk and in the lateral and posterior marginal series. For material from several rivers, the range of length of metamorphosing and macrophthalmia stages of L. fluviatilis is83–119 mm (mean 99.3 mm). Weights varied from 0.71-2.5 g (mean 1.51 g). Regression coefficients for weight on length are much lower in the macrophthalmia of L. fluviatilis than in comparable stages of L. planeri. Total oocyte counts on macrophthalmia of L. fluviatilis gave values from8000–20,000 which are in general agreement with egg counts for adults of this species in the river Severn. Observations on the earliest metamorphosing forms have shown that it is not possible at this stage to distinguish the males of the two species by the structure of the testes. The development of the lumen in the adult foregut of L. fluviatilis has been shown to be variable and in some instances the gut does not become patent until early Spring. The possibility has also been raised that, in exceptional cases, a temporary lumen may also be present for a short period in L. planeri. Observations on the foregut and dentition, together with field data, suggest that the climax of downstream migration and the onset of parastic feeding takes place in late March or April.     

Population genetics of a cyclostome species pair, river lamprey (Lampetra fluviatilis L.) and brook lamprey (Lampetra planeri Bloch)

Horizontal agarose gel electrophoresis of 24 allozyme loci in four species of Central European lampreys (321 Lampetra planeri , 83 L. fluviatilis , 11 Eudontomyzon mariae and nine Petromyzon marinus ) was used to study the 'paired species' L. fluviatilis and L. planeri . The genetic differentiation of the anadromous river lamprey ( L. fluviatilis ) from the stationary brook lamprey ( L. planeri ) was within the range of ingroup differentiation of the latter, but L. fluviatilis exhibited much greater population cohesion over a more extended geographic range: G _ST = 0.0537 versus G _ST = 0.3398, N _em = 4.402 versus N _em = 0.4856, mean genetic among-stock distances D = 0.0047 versus D = 0.0257. L. planeri populations coexisting geographically with L. fluviatilis in the Rhine and Elbe river systems were genetically more cohesive than L. planeri stocks from the Danubian basin where L. fluviatilis is absent. Danubian L. planeri populations exhibit a lower degree of heterozygosity than brook lampreys from the Rhine river system, but comprise deeper genetic lineages ( G _ST = 0.4629 versus G _ST = 0.2434), despite being sampled from a much more restricted area. Isolation-by-distance is observed for L. planeri from the Danubian but not from the Atlantic drainage basins. Transspecific gene flow between L. planeri from Atlantic drainage basins and the long-distant migrating L. fluviatilis is inferred, raising doubt on the validity of two separate biospecies. E. mariae and P. marinus are clearly differentiated from Lampetra spp. at several allozyme loci.     

Changes in haemopoietic sites during the metamorphosis of the lampreys Lampetra fluviatilis and Lampetra planeri

An investigation has been made in both the parasitic lamprey Lampetra fluviatilis and in its non-parasitic derivative Lampetra planeri of the rate at which the fat column replaces the typhlosole and nephric fold as the principal site of haemopoiesis. In the typhlosole, blood cell formation started to decline prior to the onset of external metamorphosis and had ceased within four weeks of the commencement of transformation. In the nephric fold haemopoiesis continued for several weeks in the region where the larval opisthonephros persisted but was never observed in the newly developing adult kidney. Soon after the onset of external metamorphosis the fat column started to become haemopoietic and later became the main site of blood cell formation. The rate at which the haemopoietic function was transferred from the nephric fold and typhlosole to the fat column was greater in L. fluviatilis than in L. planeri. Since a similar more rapid change in L. fluviatilis has also been found in the switch from larval to adult haemoglobin, the former type of haemoglobins may be produced only in erythrocytes originating in the nephric fold and typhlosole, whereas the latter type may be restricted to cells developed in the fat column. It is also suggested that the functional significance of the alteration in haemopoietic sites is related to changes at metamorphosis in the three regions where blood cell formation occurs.     

Metamorphosis in the paired species of lampreys, Lampetra fluviatilis (L.) and Lampetra planeri (Bloch): 2. Quantitative data for body proportions, weights, lengths and sex ratios

Representatives of the various Stages (1–9) in the metamorphosis and short adult life of the brook lamprey, Lampetra planeri , were used to provide quantitative data to describe the pattern of changes that take place during this period of the life cycle. During the transformation of the oral hood into an oral disc, this ventral region of the snout first declines in length before increasing markedly. The branchial region declines in length and remains relatively smaller than in the ammocoete, a feature probably correlated with the concomitant internal changes in the pharynx which permit a change from a unidirectional to a tidal respiratory flow. The eye increases in size only through the initial Stages (1–5). The main period of fin enlargement occurs rather later in metamorphosis than the above changes.
During maturation the two initially separate dorsal fins increase further in height and eventually become separated only by a notch. Differences between the two sexes are found in the relative size of the disc and fins, the larger structures of the male probably being related to their greater activity in spawning behaviour. Measurements on metamorphosed L. fluviatilis confirm that in this species the eye and disc are relatively larger than in L. planeri. Data are presented which illustrate that in collections taken during the metamorphosis of both species the males are smaller and slightly more numerous than the females and that transformation is initiated at a longer length in L. planeri than in L. fluviatilis .     

A multivariate morphometric and meristic description of a population of freshwater-feeding river lampreys, Lampetra fluviatilis (L.), from Loch Lomond, Scotland

Adults of a freshwater-feeding population of Lampetra fluviatilis (L.) in Loch Lomond, Scotland, were trapped on their upstream (spawning) migration in the main inflow to the loch, the Endrick Water. These animals could be distinguished from the more typical anadromous upstream migrant L. fluviatilis , from the Endrick Water and the nearby catchment of the River Forth, principally by their smaller total length, greater disc, snout and eye size and much darker pigmentation. These, and other morphometric and meristic characteristics have been compared for populations of L. fluviatilis and Lampetra planeri (Bloch) from the Lomond and Forth catchments using univariate and multivariate statistics. The Endrick Water 'dwarf' L. fluviatilis was found to differ significantly from the other Lampetra studied, in a number of body proportions in addition to those traits mentioned above, and also in mean numbers of trunk myomeres and teeth. These differences are likely to be due to a combination of the effects of genetic isolation, and the short (freshwater) feeding phase of the 'dwarf' L. fluviatilis , which may, therefore, represent the intermediate stage between parasitic and non-parasitic forms in the L. fluviatilis/L. planeri species pair.     

The systematics of British larval lampreys

Several taxonomic characters, including the number of trunk myomeres, proportional body measurements and patterns of external and internal pigmentation, have been investigated in populations of ammocoetes belonging to the three species of lampreys found in British rivers. Petromyzon marinus can be readily distinguished by the pigmentary pattern in the tail and on the oral hood, and by the much higher number of trunk myomeres. Although no character can be used to separate with certainty the larvae of Lampetra fluviatilis and Lampetra planeri , significant differences were found between the mean number of trunk myomeres in the two species. Moreover, the values for ammocoetes were virtually identical to those of the respective post-larval stages, at which phase in the life cycle the two Lampetra species are clearly distinguishable. The incidence of pigmentation on the tongue precursor was much higher in larval populations of L. fluviatilis than in those of L. planeri and there was also a difference in the rate of change in the relative length of the trunk and tail. Where comparisons can be made, the results in this paper are in agreement with those of most other European and North American workers. The data differ markedly, however, from that contained in the only other previous detailed systematic study on British larval lampreys, which was subsequently used as the basis for taxonomic keys.     

Brown trout Salmo trutta redd superimposition by spawning Lampetra species in a lowland stream

Reproductive interaction between sympatric lampreys and salmonids was studied. The superimposition of brown trout Salmo trutta redds by spring-spawning river lamprey Lampetra fluviatilis and brook lamprey Lampetra planeri was examined in a small lowland stream of western Lithuania. A high superimposition rate of S. trutta redds by both L. fluviatilis (up to 83%) and L. planeri (up to 48%) was found, when the spawning intensity of Lampetra spp. was high. The occurrence of this phenomenon is the result of the overlap in the spawning habitat preferences at the reach-scale and at the microhabitat scale for the three species. One of the main requirements for Lampetra spp. spawning site selection was the negative streambed slope, an essential trait of the pool-riffle transitional zone. The structure of the salmonid redd created a considerable negative microhabitat slope suitable for Lampetra spp. spawning, which put the redds under a higher susceptibility to be superimposed. The timing of Lampetra spp. spawning overlapped closely with the emergence of S. trutta fry, suggesting a probable ecological effect of superimposition on S. trutta in the pre-emergent and emerging stages.     

Intraspecific developmental variation in the life cycle of the Andean Treefrog (Boana riojana): A temporal analysis

Intraspecific variation during the anuran larval period has been analyzed mainly in relation to the timing of metamorphosis and body size at metamorphosis. However, other traits may vary as well. We examined two developmental series of Boana riojana from the same population in two consecutive years and describe intraspecific variation in larvae of this species. We discuss how variation, if present, may influence its life cycle. We found that both larval series differed in the larval period length, one twice as long as the other. This variation primarily depended on when breeding occurred, metamorphosis was achieved during late spring in both generations and at similar sizes, and only the rate of larval development during premetamorphosis varied extensively between years. This is consistent with thyroid gland activity because when it became active the developmental trajectory became more canalized. No variation of staging sequence occurred in relation to the different durations of the larval period. However, in the long-lasting series we found two different morphs. Also, integument, thyroid gland, skeleton, and testis differentiation events occurred at the same developing stages. In contrast, ovarian differentiation proceeded at the same absolute age in both series. Sexual dimorphism becomes evident within the year after metamorphosis. The intraspecific heterochrony that we describe for the larval development of B. riojana does not lead to phenotypic variation at the end of metamorphosis. We discuss the importance of analyzing growth and development independently. Each proceeds differently in time, but with an interdependence at some point, because size and shape do not vary at the end of metamorphosis.     

Contribution to the knowledge of the biology of Cyclostomes: activity of aldolase in Lampetra planeri (Bloch) during its biological cycle

It has been ascertained that in the Lampetra planeri (Bloch) the muscle and liver aldolase level in the last larval period in which lipids are accumulated strongly increases; this may be due to an increased synthesis or diminished degradation of the enzyme.     

The critical period for wing pattern induction in the polyphenic tropical butterfly Bicyclus anynana (Satyrinae)

Adults of the butterfly Bicyclus anynana express striking phenotypic plasticity. A wet season form has conspicuous marginal eyespots and a medial pale band which are much reduced in the dry season form. These alternative forms are produced after rearing at high or low temperatures, respectively. We used 'window' experiments involving switching of larvae and pupae between high and low temperatures at different stages during development to examine the timing of sensitivity to environmental temperature. The final, fifth larval instar is shown to be especially sensitive. The fourth larval instar and the very early pupal period are also sensitive. It is argued that an increasing sensitivity during growth is ecologically adaptive since the late larval environment will be the most accurate predictor for the adult environment in which the wing phenotype is subject to selection. The period of sensitivity is not as short as a few days. This may minimize the chance of any 'mistakes' in matching the adult phenotype to the season because of short-term environmental fluctuations during the larval period. The observed sensitivity occurs as late as possible during growth since the wing pattern is developmentally determined at the end of the early part of the pupal stage.     

Evolutionary history of lamprey paired species Lampetra fluviatilis (L.) and Lampetra planeri (Bloch) as inferred from mitochondrial DNA variation

A remarkable trend in the evolution of lampreys is the occurrence in most genera of 'paired species', in which the parasitic anadromous lampreys are believed to have given rise to nonparasitic freshwater resident populations. The present work examines the phylogeography of the European paired species Lampetra fluviatilis and Lampetra planeri, in an attempt to elucidate species pair evolutionary history. We studied sequence variation in cytochrome b and ATPase 6, 8 mitochondrial genes in 63 individuals from 21 localities of the paired species throughout their distribution range. Results from the phylogenetic and nested clade analyses were largely consistent, suggesting the existence of three major evolutionary lineages: lineage I and possibly lineage II are widespread throughout Europe, while the most ancestral lineage III is apparently restricted to the Iberian Peninsula. The high genetic diversity observed in the Iberian Peninsula is probably the result of refugial persistence and subsequent accumulation of variation over several ice ages, whereas the low levels of genetic diversity observed in central and northern Europe should reflect a rapid postglacial colonization. Results suggest that L. planeri originated within at least two distinct evolutionary lineages, rejecting the single origin hypothesis. The observed lack of taxa monophyly within lineage I may be the result of ongoing gene flow if the two taxa are alternate life-history forms of a single species. However, structure within lineage I is also consistent with the hypothesis of divergence of taxa after postglacial dispersion (around 2000 generations ago) with incomplete lineage sorting. Further testing of the alternative hypotheses is warranted.     

Gill Morphometrics of the Lampreys Lampetra fluviatilis (L.) and Lampetra planeri (Bloch)

Morphometric measurements have been made on various gill components of different stages in the life cycle of the anadromous parasitic lamprey, Lampetra fluviatilis, and its nonparasitic derivative Lampetra planeri. The total gill area, expressed in terms of body weight, of both larval (1462–2717 mm² g^–1) and adult (1402–2337 mm² g^–1) L. fluviatilis are greater than those previously recorded in the rather meagre literature on lamprey gill measurements and are comparable with those found in the most active teleosts. The gills of the two Lampetra species are apparently identical in the larval stages and those of metamorphosing and adult L. planeri are similar to those of metamorphosing L. fluviatilis. Although the pharyngeal arrangement of lampreys differs greatly from that of teleosts, there are many features of the gills indicative of convergence between the two groups. Thus, in a given stage in the life cycle of lampreys, the secondary lamellae on either side of the filaments also alternate, become more widely spaced as the filament length increases and increase in area as the body weight becomes greater. Furthermore, the fractional cumulative increase in secondary lamellae area along a line following the presumed direction of water flow is also represented by a sigmoid curve. While at metamorphosis the pharynx becomes considerably modified to accommodate the change from a unidirectional to a tidal respiratory water flow, the total gill areas of the ammocoete are similar to those of metamorphosing stages which have attained adult characteristics. However, there are clearly differences in some of the components that influence and contribute towards the total gill area. Thus, in terms of body weight, the number and total length of the filaments and the total number of secondary lamellae, together with the number of secondary lamellae found on a given distance of filament, are greater in late metamorphosing stages, while the reverse is true for the average bilateral area of the secondary lamellae which is considerably greater in ammocoetes.     

Shifts in the microhabitat of larval and juvenile roach, Rutilus rutilus (L.), in a floodplain channel

The microhabitat of roach larvae (stages L1–L6) and 0+ juveniles was examined in a braided channel (Ilon) of the Upper Rhone River, France, from weekly samples (spring-autumn 1985) using Point Abundance Sampling by electrofishing. During early development, microhabitat exploitation was characterized by two transitions: the first at larval step L3, when morphological and physiological attributes permit the active choice of microhabitat; the second at the end of the larval and start of the juvenile period. At larval step L3, and up to L5, roach were strongly associated with lentic waters of medium depth (0.5–1.0m) and with either macrophytes or ligneous debris. The L6 and 0+ juveniles were found more often than expected in shallower waters (< 0.2m, 0.2–0.5 m) and were no longer associated with dense vegetation or ligneous debris. However, both larval (L3–L6) and 0+ juvenile roach preferred silty sand and/or silted gravel, and a lack of current. This shift, from moderately deep waters with vegetation and/or branches as larvae to open shallow waters as juveniles, demonstrates the young roach's perception and response to environmental change, with shallows exploited as a refuge from predation when protective structures become scarce or unavailable.     

Heart Ratios at Different Stages in the Life Cycle of Lampreys

Heart ratios (heart weight/body weight. ×100) have been obtained for freshly killed and preserved (10% formalin) specimens of the lampreys Petromyzon marinus, Lampetra fluviatilis and L. planeri. Values for comparable stages were significantly higher in the preserved material, probably as a result of retention and fixation of blood in the heart chambers. The regression slope for the logarithmic relationship between heart and body weights was close to one in all stages of the three species investigated. A marked increase was observed in the heart ratio during metamorphosis, apparently attributable to an increase in the number and density of myocardial fibres. Heart ratios in the lamprey were generally higher than for most other fishes, with values of 0.1 and 0.3 being obtained for example in the case of freshly killed ammocoete and sexually mature adult stages of L. fluviatilis.     

Allozyme Polymorphisms and Biochemical-Genetic Taxon Markers in Lampreys (Lampetra, Eudontomyzon, Petromyzon)

Variation at 23 putative enzyme-coding loci was scored in 424 lampreys, including 321 European brook lampreys (Lampetra planeri), 83 European river lampreys (L. fluviatilis), 11 Ukrainian brook lampreys (Eudontomyzon mariae), and nine sea lampreys (Petromyzon marinus). Twelve polymorphisms are described for Lampetra species (LDH*, SOD-2*, PNP*, AAT-1*, AK-1*, ES-2*, LGL*, MPI*, GPI-1*, GPI-2*, PGM*, IDHP-2*), and two each for E. mariae (GPI-1*, ME-2*) and P. marinus (MDH-1*, ME-2*). Diagnostic allozymes are presented for the discrimination of lamprey taxa, some of which are difficult to recognize from the morphology of ammocoetes larvae, the life stage usually encountered when collecting cyclostomes. The allelic markers described permit the clear allocation to a genus, except for the species L. fluviatilis and L. planeri, which are not differentiated by qualitative allozyme analysis.     

Heated competition: how climate change will affect non-native pumpkinseed Lepomis gibbosus and native perch Perca fluviatilis interactions in the U.K

Heated and ambient temperature experimental ponds were used to examine competition between introduced pumpkinseed Lepomis gibbosus and native perch Perca fluviatilis in England, U.K., and how these interactions are likely to change under climate warming conditions. Results from three sets of two-week experiments indicated that in both species and in all sets, growth was faster in heated than in ambient temperature ponds. Growth of both P. fluviatilis and L. gibbosus in sympatry did not differ significantly from that observed in allopatric ponds. Diet analysis indicated that increased resource partitioning occurred when P. fluviatilis and L. gibbosus were reared in sympatry, with P. fluviatilis shifting to a diet higher in microcrustaceans. The results do not support the previous claims of adverse effects of L. gibbosus on P. fluviatilis populations. Under conditions of climate change, however, which have been demonstrated experimentally to enhance L. gibbosus recruitment, this species is expected to become invasive in England, resulting in higher densities that may exert a stronger competitive effect than examined in this study.     

Photoperiodic induction of the pupal diapause in the tobacco hornworm, Manduca sexta

A study was made of photoperiodic induction of the facultative pupal diapause in the tobacco hornworm, Manduca sexta, reared on artificial diet in the laboratory. The species entered a prolonged diapause when the egg and larval feeding stages were reared in daily photoperiods of 13·5 hr or less. Diapause was induced in all insects at photoperiods ranging from 1 to 13 hr, and part of the population entered diapause at only 15 to 30 min of light per day. Photoperiods of 14 hr or more and continous darkness prevented diapause. Duration of diapause varied with the inductive photoperiod in which the hornworms were reared during the sensitive period. Insects reared in longer diapause-inducing photoperiods within a range of 12 to 13·25 hr remained in diapause longer than those reared in shorter photoperiods. There was no difference in the rate of larval development of hornworms reared in diapause-inducing vs diapause-preventing photoperiods. Temperatures of 26 to 30°C were most favourable for the photoperiodic induction of diapause; at 21°C, the critical photoperiod and incidence of diapause were decreased. Diapause induction was suppressed by low (18°C) and higher (33°C) temperatures. The number of inductive 12L:12D (light = 12 hr; dark = 12 hr) cycles required to induce diapause ranged from as few as 5 for some insects to as many as 12 for others when the post-inductive régimen was continuous light, but with insects previously held in continuous dark, as few as 2 12L:12D cycles during the last 2 days of larval feeding induced diapause in 38 per cent of the population. Only 3 to 4 cycles of 15L:9D during the final larval instar reversed inductive effects of 14 to 15 12L:12D cycles. Photoperiodic sensitivity extended from the late embryo to the end of larval feeding but showed considerable fluctuation during development with maximum sensitivity occurring just before egg hatch and during larval growth.Light breaks applied at different times during the dark period of 12L:12D cycles generated different response curves, depending on the number of cycles in which light breaks were repeated. When repeated for 6 cycles, a unimodal response curve was obtained; 10 cycles produced a bimodal curve and light breaks given for 18 cycles throughout the sensitive period averted diapause regardless of time of night applied. It is suggested that diapause is regulated by a photo- and thermolabile substance that accumulates during long nights (11 hr or more) and acts during the early pupal stage to inhibit the translocation and release of development-promoting neurosecretion from the brain.     

Characteristics of the Norwegian Coastal Current during Years with High Recruitment of Norwegian Spring Spawning Herring (Clupea harengus L.)

Norwegian Spring Spawning herring (NSSH) Clupea harengus L. spawn on coastal banks along the west coast of Norway. The larvae are generally transported northward in the Norwegian Coastal Current (NCC) with many individuals utilizing nursery grounds in the Barents Sea. The recruitment to this stock is highly variable with a few years having exceptionally good recruitment. The principal causes of recruitment variability of this herring population have been elusive. Here we undertake an event analysis using data between 1948 and 2010 to gain insight into the physical conditions in the NCC that coincide with years of high recruitment. In contrast to a typical year when northerly upwelling winds are prominent during spring, the years with high recruitment coincide with predominantly southwesterly winds and weak upwelling in spring and summer, which lead to an enhanced northward coastal current during the larval drift period. Also in most peak recruitment years, low-salinity anomalies are observed to propagate northward during the spring and summer. It is suggested that consistent southwesterly (downwelling) winds and propagating low-salinity anomalies, both leading to an enhanced northward transport of larvae, are important factors for elevated recruitment. At the same time, these conditions stabilize the coastal waters, possibly leading to enhanced production and improved feeding potential along the drift route to Barents Sea. Further studies on the drivers of early life history mortality can now be undertaken with a better understanding of the physical conditions that prevail during years when elevated recruitment occurs in this herring stock.     

Growth, dynamics, and distribution of a population of the brook lamprey Lampetra planeri in a South Swedish stream

Size and structure of a South Swedish population of Lampetra planeri larvae were studied during 5 yr. Population size fluctuated considerably within the studied section of stream with a general decline. The fluctuations were positively and strongly correlated (multiple R²= 0.85) with the population density at the beginning of each analysed period, while poorly correlated with such variables as heavy precipitation and temperature. Several factors suggest the entire larval population drifts downstream: fluctuations in the studied stream section, increasing mean size downstream, and incongruity in longitudinal distribution in surveys, performed with a 5-yr interval. Growth, estimated both in caged larvae and from length frequency data, occurred mainly in autumn. High density in cages reduced the growth rate of larvae.