Evidence of a recent decline in river lamprey Lampetra fluviatilis parasitism of a nationally rare whitefish Coregonus lavaretus: is there a diamond in the ruffe Gymnocephalus cernuus?

Lamprey‐induced scarring of the nationally rare Coregonus lavaretus, a known host of a freshwater‐resident population of European river lamprey Lampetra fluviatilis, was found to have declined precipitously since the establishment of several non‐native fishes in Loch Lomond. Evidence presented in this study points to the possibility that L. fluviatilis in this lake may have altered its trophic ecology in response to the negative impact that non‐native species, in particular ruffe Gymnocephalus cernuus, have had on their favoured host.     

River lamprey present an unusual predation threat to Atlantic salmon smolts in Lough Neagh,Northern Ireland

A new monitoring programme on the Lough Neagh catchment has documented a high incidence of river lamprey, Lampetra fluviatilis L., predation on Atlantic salmon smolts, Salmo salar L. In total 470 smolts were examined during the 2020 emigration period with 168 fish (36%) exhibiting lamprey scars of which 57 were lightly scarred and 111 were classed as heavily scarred. Lamprey predation was not size selective on Lough Neagh S. salar smolts.     

Has habitat heterogeneity promoted phenotypic and ecological sub‐structuring among a Coregonus lavaretus population in a large Scottish lake?

Differences in stable‐isotope values, morphology and ecology in whitefish Coregonus lavaretus were investigated between the three basins of Loch Lomond. The results are discussed with reference to a genetic investigation to elucidate any substructuring or spawning site fidelity. Foraging fidelity between basins of Loch Lomond was indicated by δ¹³C and δ¹⁵N values of C. lavaretus muscle tissue. There was, however, no evidence of the existence of sympatric morphs in the C. lavaretus population. A previous report of two C. lavaretus‘species’ in Loch Lomond probably reflects natural variation between individuals within a single mixed population.     

Morphological abnormalities in a population of Lampetra planeri,with a short review of petromyzontid teratologies

Reported here are several deviations (n = 15; 1·96%) from typical morphology in a large sample (n = 767) of European brook lamprey Lampetra planeri from a single population in the Loch Lomond catchment; this includes one specimen bearing a true anal fin. A brief review of petromyzontid teratology is provided.     

The fish community of Loch Lomond,Scotland: its history and rapidly changing status

The fish community of Loch Lomond is of national importance. Its diversity of species and rare populations of powan (Coregonus lavaretus) and freshwater feeding river lampreys (Lampetra fluviatilis) warrant high conservation status. It is also of value for its sport fisheries for sea-trout (Salmo trutta), salmon (Salmo salar) and pike (Esox lucius). Historical records demonstrate that the species composition of the fish community has remained stable over a very long period until recently when a series of introductions of fish species new to the catchment has resulted in successful colonisation by a number of species. These have resulted in fundamental changes in the ecosystem. Here using historical records the long-term stability of the fish community is examined, recent rapid changes in the fish community are documented and some of the resultant effects of changes in the fish community are demonstrated.     

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.     

Genetic diversity,endemism and phylogeny of lampreys within the genus Lampetra sensu stricto (Petromyzontiformes: Petromyzontidae) in western North America

Phylogenetic structure of four Lampetra species from the Pacific drainage of North America (western brook lamprey Lampetra richardsoni, Pacific brook lamprey Lampetra pacifica, river lamprey Lampetra ayresii and Kern brook lamprey Lampetra hubbsi) and unidentified Lampetra specimens (referred to as Lampetra sp.) from 36 locations was estimated using the mitochondrial cytochrome b gene. Maximum parsimony and Bayesian inferences did not correspond with any taxonomic scheme proposed to date. Rather, although L. richardsoni (from Alaska to California) and L. ayresii (from British Columbia to California) together constituted a well‐supported clade distinct from several genetically divergent Lampetra populations in Oregon and California, these two species were not reciprocally monophyletic. The genetically divergent populations included L. pacifica (from the Columbia River basin) and L. hubbsi (from the Kern River basin) and four Lampetra sp. populations in Oregon (Siuslaw River and Fourmile Creek) and California (Kelsey and Mark West Creeks). These four Lampetra sp. populations showed genetic divergence between 2·3 and 5·7% from any known species (and up to 8·0% from each other), and may represent morphologically cryptic and thus previously undescribed species. A fifth population (from Paynes Creek, California) may represent a range extension of L. hubbsi into the Upper Sacramento River.     

Life histories of the powan,Coregonus lavaretus (L.) (Salmonidae,Coregoninae) of Loch Lomond and Loch Eck

Four aspects of the life histories of the two populations of powan Coregonus lavaretus (L.) in Scotland are described: growth (Eck powan are shorter and with greater year to year variance than Lomond); sexual maturation (Eck powan mature younger, but at similar weight to Lomond); spawning (timing in Eck varies, but is consistent in Lomond); and recruitment/mortality (fecundity, sex ratios, and mortality also vary in the short term). Short term differences between the physiological ecology of the populations can be ascribed to the size and topography of the lochs. Long term differences are more difficult to account for, and are more important in that they may signal changes in sustainability. Conservation of powan must be considered in terms of their synecological relationships, not in isolation.     

Incidence of lamprey marks on Lake Sturgeon (Acipenser fulvescens Rafinesque, 1817) in the St. Clair – Detroit River System: Implications for Sea Lamprey (Petromyzon marinus Linnaeus, 1758) effects

Laurentian Great Lakes Lake Sturgeon (Acipenser fulvescens) are hosts to lamprey species, including native Silver Lamprey (Ichthyomyzon unicuspis) and invasive Sea Lamprey (Petromyzon marinus). Silver Lamprey coevolved with Lake Sturgeon and cause negligible mortality, but Sea Lamprey can negatively affect Lake Sturgeon populations. Sea Lamprey abundance in Lake Erie has been above targets set by resource managers, with the St. Clair – Detroit River System (SCDRS) suspected as a source of Sea Lamprey production into Lake Erie. This study summarizes lamprey marking on Lake Sturgeon captured during agency assessment surveys in the SCDRS since 1996 and provides insight on the potential for Sea Lamprey to negatively affect Lake Sturgeon in the SCDRS. Lamprey marks (any lamprey species) were noted on 48.2% of Lake Sturgeon (2.5 marks/fish) and 3.3% of Lake Sturgeon assumed to be susceptible to mortality by Sea Lamprey (<760 mm TL; 0.06 marks/fish). Silver Lamprey were the only lamprey species found attached to Lake Sturgeon and there was no difference between oral disc diameters of Silver Lamprey and marks measured on Lake Sturgeon in Lake St. Clair and the lower St. Clair River (p = .45). Based on logistic regression, probability of at least one lamprey mark increased with Lake Sturgeon total length and was highest in Lake St. Clair. The probability of observing at least one lamprey mark on a 760 mm Lake Sturgeon was 8.1% or less for each sampling location in the SCDRS aside from Lake St. Clair (28.1%). Results suggest that parasitism of Lake Sturgeon by Sea Lamprey in the SCDRS is rare, particularly for Lake Sturgeon <760 mm TL. Low incidence of lamprey marks on Lake Sturgeon assumed to be susceptible to mortality from Sea Lamprey parasitism and zero occurrence of Sea Lamprey being observed attached to a Lake Sturgeon suggest Sea Lamprey at their current abundance likely have little effect on the Lake Sturgeon population in the SCDRS. Caution should be taken when using mark size to assign marks to lamprey species as there is substantial overlap among species oral disc diameters, potentially inflating the perceived impact of Sea Lamprey on Lake Sturgeon in areas with native lampreys.     

Phylogeography of the European brook lamprey (Lampetra planeri) and the European river lamprey (Lampetra fluviatilis) species pair based on mitochondrial data

The European river lamprey Lampetra fluviatilis and the European brook lamprey Lampetra planeri (Block 1784) are classified as a paired species, characterized by notably different life histories but morphological similarities. Previous work has further shown limited genetic differentiation between these two species at the mitochondrial DNA level. Here, we expand on this previous work, which focused on lamprey species from the Iberian Peninsula in the south and mainland Europe in the north, by sequencing three mitochondrial marker regions of Lampetra individuals from five river systems in Ireland and five in southern Italy. Our results corroborate the previously identified pattern of genetic diversity for the species pair. We also show significant genetic differentiation between Irish and mainland European lamprey populations, suggesting another ichthyogeographic district distinct from those previously defined. Finally, our results stress the importance of southern Italian L. planeri populations, which maintain several private alleles and notable genetic diversity.     

Freshwater and wetland plant communities of Loch Lomond

Information on the aquatic and wetland vegetation of Loch Lomond, Scotland, is summarised for the period 1957–90. Aquatic macrophyte growth is estimated to occupy about 1% of the loch's total surface area, being limited to the 0–10 m euphotic zone, and probably excluded from much of this area by wave disturbance and unsuitable substrates. Aquatic vegetation is however abundant in sheltered bays and less-exposed shorelines, particularly in the South Basin of the loch. Although Littorella uniflora (L.) Aschers is the commonest, ubiquitous, submerged plant of the loch, three separate euhydrophyte communities have been identified within the loch. One of these is characterised by abundant stands of Elodea canadensis Michx., which appears to have invaded Loch Lomond some time between 1967–88, and has rapidly spread throughout the loch.     

Reversible metabolic depression in lamprey hepatocytes during prespawning migration: dynamics of mitochondrial membrane potential

The lamprey (Lampetra fluviatilis L.) is an extant representative of the ancient vertebrate group of Agnathans. During the prespawning migration (the river period of life from autumn until spring) lamprey hepatocytes exhibit widely different energy states: a high-energy state in autumn and spring, corresponding to a normal physiological standard, and a low-energy state in winter, which is provoked by prolonged starvation and profound metabolic arrest. In spring the restoration of energy status (return to an active state) is associated with hormonally induced lipolysis of the lipid droplets stored in the cells. Lamprey hepatocytes demonstrate an aerobic metabolism based on oxidation of free fatty acids. The dynamics of mitochondrial membrane potential (MMP) were measured throughout the prespawning migration. Pharmacological inhibition of the electron transport chain decreased the MMP and caused extensive depletion of cellular ATP without loss of cell viability. The potential molecular mechanisms responsible for winter metabolic depression in lamprey hepatocytes are discussed.     

Zooplankton in Loch Lomond: perspectives,predation and powan

Published and unpublished accounts of the zooplankton community of Loch Lomond are reviewed. Loch Lomond's zooplankton community is unique amongst those of Scotland's other large lochs. Holopedium gibberum appeared in Loch Lomond in the years between the studies of Slack & Hamilton in the 1950s and zooplankton surveys in the late 1970s.The annual cycles of Eudiaptomus gracilis, Cyclops strenuus abyssorum and Mesocyclops leuckarti are described for the mid basin of Loch Lomond. E. gracilis overwinter mainly as adult with variable egg production during their lifespan of 5–8 months. The cyclopoid copepods have variable life history characteristics. C. str. abyssorum produce a single generation per year, overwinter by means of resting eggs and a small population of planktonic adults, while M. leuckarti is present in the plankton throughout the year, but in low numbers during winter.Zooplankton populations in Loch Lomond are predated by the diverse fish community which includes a facultative planktivore in the powan, Coregonus lavaretus. Powan feed heavily on zooplankton from late spring until late autumn. Powan display selectivity in the prey species taken, in the size distribution and morph types taken within prey species. Zooplankton species, morphs and forms in Loch Lomond tend to reflect this predation.Zoology Department, University of St. AndrewsZoology Department, University of St. Andrews     

Lethal Temperatures in Ammocoetes of Four Species of Lampreys

Abstract An investigation has been made of the resistance time and upper lethal temperature of ammocoetes of four species of lampreys provided with a substrate into which they could readily burrow. In general, ammocoetes burrowed after transfer from the acclimation to the experimental temperature baths and later came out of the substrate only in lethal temperatures. A relationship was observed between the resistance time and the time taken to emerge, with the resistance time increasing exponentially with decreasing experimental temperature. In Ichthyomyzon fossor, landlocked Petromyzon marinus, Lampetra (Lethenteron) Lamottenii and in Lampetra (Lampetra) planeri from two different times of the year, the incipient lethal levels over a two week experimental period for larvae acclimated to 15° G were respectively 30.5, 30, 29.5, 28.5 and 28° C. Values for P. marinus acclimated to 5 and 25° C were respectively 29.5 and 31° C, whereas in L. planeri they were 28 and 29° C in April/May and 27 and 29° C in July/August. Extrapolation of the results for the three acclimation temperatures yielded ultimate incipient lethal levels of 31.4° G in P. marinus and 29.2 and 29.4° C for L. planeri examined in the spring and summer respectively.     

The genus Lessonia Bory (Phaeophyta,Laminariales) in Southern Chile and Argentina

Study of Lessonia in southern Chile and Argentina indicates that it is represented there by three species: L. nigrescens Bory, L. flavicans Bory and L. vadosa Searles sp. nov. Plants of these three species have been confused with each other in the literature, but appear clearly separable. Lessonia nigrescens is distinguished from the others by its massive, solid holdfast and numerous stipes. In Lessonia flavicans and L. vadosa a single stipe arises from a branched holdfast. These two species are distinguishable from each other in details of blade morphology, anatomy, and ecology. The taxonomic status of plants in South America outside of the study area remains unresolved. Populations in the Falkland Islands and in northern Chile in particular are in need of further study.     

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.     

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.     

Alternative migration and host parasitism strategies and their long-term stability in river lampreys from the River Endrick, Scotland

The stability of a discrete body size dimorphism of sexually mature river lamprey Lampetra fluviatilis from the River Endrick, Scotland, was examined over a 21 year period. Stable isotope analysis was used to test the hypothesis that the two size forms comprise individuals with differing migration and parasitic foraging strategies. Maturing river lamprey and the brook lamprey Lampetra planeri were trapped over 3 months each year in the periods 1983–1984 and 2004–2005. Brook lamprey catches and catches of both species combined showed no significant trend in catch rate with time. The catch rate of small body size river lamprey declined between 1983–1984 and 2004–2005 (although the difference did not reach statistical significance; P = 0·055). In contrast, there was a significant increase in the catch rate of the large body size river lamprey and as a consequence, a significant change in the relative proportion of each of the two river lamprey morphs over the study period. Analysis of the stable isotopes of C and N in muscle tissue showed that brook lamprey tissue derived its carbon from a freshwater source and had a δ¹³C more consistent with that of the River Endrick than with Loch Lomond. δ¹⁵N values for this species showed it to be feeding at the base of the food chain, consistent with filter feeding as an ammocoete. The large body size and the small body size river lamprey adults differed substantially in their δ¹³C values, with the small body size δ¹³C signature indicative of a freshwater carbon source and the large body size morph of a marine source. The small body size morph had a δ¹³C signature that was consistent with that of Loch Lomond powan Coregonus lavaretus suggesting that they share a common carbon source. The large body size morph was clearly feeding at a higher trophic level than the small body size morph. A single small body size river lamprey individual with typical morphology for that group, however, had C and N signatures that clustered with those of the large body size morphs. This individual had either migrated to sea to forage, as is typical for the species, or had been feeding on an anadromous fish with a strong marine C signature in fresh water. It is concluded that the body size dimorphism is indicative of a differential migration and foraging strategy in the parasitic phase of the life cycle of river lamprey at this site.     

Post‐zygotic hybrid viability in sympatric species pairs: a case study from European lampreys

Ecological speciation mechanisms are widely assumed to play an important role in the early stages of divergence between incipient species, and this especially true of fishes. In the present study, we tested explicitly for post‐zygotic barriers to gene flow between a sympatric, recently diverged lamprey species pair that likely arose through ecological divergence. Experimental in vitro hybridization between anadromous parasitic Lampetra fluviatilis and resident nonparasitic Lampetra planeri resulted in a high proportion of embryos capable of attaining the burrowing pro‐larval stage, strongly indicating no post‐zygotic barriers to gene flow between these species. A sympatric, locally‐adapted resident parasitic form of L. fluviatilis was also found to successfully hybridize with both members of this species pair. The consequences of these findings are discussed in the context of lamprey speciation. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 108 , 378–383.     

Three new species of Lysipomia (Lobeliaceae) endemic to the páramos of southern Ecuador

Three species of Lysipomia, L. caespitosa, L. cylindrocarpa, and L. speciosa, from a tropical alpine ecosystem called páramo, are described, illustrated, and compared to related species. Notes on morphology, especially floral presentation, and general ecology are given where possible. A published key to the species in Ecuador is revised to include the three new species plus L. multiflora McVaugh, a species previously known only from northern Peru.