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The extended freshwater life of Petromyzontiformes larvae (ammocoetes) takes place mainly in lotic waters where it has been studied in detail. Information on ammocoetes in lentic habitats is extremely limited, with few data on the occurrence of specimens in ponds or large lakes. The area of the Smolenskoye Poozerye National Park has a dense network of lakes and rivers, with streams abundantly populated by lamprey larvae of the genera Lampetra and Eudontomyzon. In this paper, we evaluate the habitability of lakes belonging to such systems and the manner in which they are populated by lamprey. Larval habitat identification and documentation (bed analysis where ammocoetes exist, including ground composition and biotic and abiotic habitat characteristics) were carried out, resulting in the first documentation of larvae and metamorphic specimens of Eudontomyzon sp. inhabiting littoral areas of small lakes. These specimens were found in the fringe biotopes between the river mouth and lakes. One peculiarity that distinguishes the lentic habitats from the typical river ones is the absence of distinct currents. Freshwater flows diffusely into lakes, mixing continuously at the sites we studied. Preferred habitats for Eudontomyzon sp. consisted of swales filled with debris carried from inflowing streams and deposited as current diminished. In ammocoete beds of this type, the examined larvae had a good trophic position. Such habitats can be both permanent and ephemeral as tributary water levels changes. During flood events, elevated velocity can flush debris and ammocoetes away from these fringe habitats and further into a lake. No accentuated active decomposition was found in the areas where the ammocoetes were present. The results of the study indicate that Petromyzontiformes larvae may be more common in lentic habitats than is currently documented.

The Ukrainian brook lamprey Eudontomyzon mariae is the most widespread lamprey species in eastern Europe. Although E. mariae is generally considered a derivative of Eudontomyzon danfordi, an exclusively freshwater parasitic species, it has alternatively been suggested that it was recently derived from a now extinct anadromous Black Sea ancestor. Several non-parasitic lampreys and the landlocked sea lamprey, which have recently evolved from anadromous ancestors, still develop a seawater-type mitochondria-rich cell (SW-MRC) in their gills. In contrast, this cell type is not present in the gills of either Lampetra aepyptera, a non-parasitic lamprey of ancient origin, or the parasitic Ichthyomyzon unicuspis and I. castaneus that likewise have long evolutionary histories in fresh water. Eudontomyzon mariae from the Vistula River in the Baltic River basin does not possess SW-MRC, which is inconsistent with a recent origin from an anadromous ancestor. Mitochondrial DNA sequence data were thus used to infer the relationship between different populations of E. mariae and E. danfordi, and to reconstruct the transition from anadromy to freshwater residency. The results suggest that E. mariae evolved independently in the Baltic, Black, and Caspian Sea basins, and not recently from an anadromous ancestor. Although E. mariae in the Danube River may have arisen relatively recently from E. danfordi (differing by 0.7–1.1% in cytochrome b gene sequence), other E. mariae populations (including in the Vistula River) are genetically closer (0.6%) to the hypothetical ancestor of both E. mariae and E. danfordi. That ancestor was probably a freshwater resident, since SW-MRCs are not rapidly lost following confinement in fresh water.

The stream bed is considered as an important key habitat, yet there is limited knowledge on the effects of burrowing organisms and the resulting bioturbation on this habitat. In this study, a standardized laboratory experiment was used to compare the impact of burrowing lamprey larvae (Eudontomyzon sp.) on physical and chemical habitat properties as well as microbial community composition in freshwater substrates to a control without animals. We hypothesized that the burrowing activity of the lamprey larvae increases the influx of open water into interstitial areas with consequences for oxygen availability and thus for microbial community composition. The study revealed a strong increase in oxygen availability (oxygen concentration and redox potential) and nitrate concentrations in interstitial water in the lamprey treatment, indicative of an increased exchange with open water. Over time, a shift in microbial community composition was observed in the lamprey treatment, which was dominated by aerobe bacteria. Because the functional relevance of lamprey larvae for freshwater ecosystems is evidentially high, they can clearly be considered as ecosystem engineers. Their important ecosystem functions should be recognized in conservation and management plans.

The Eudontomyzon danfordi characteristic habitats state of Maramureş Mountains Nature Park varies greatly, 19.05% are in excellent conservation status, 47.62% are in good/average status and 33.33% are in a partially degraded condition. The identified human impact categories which induced the decreasing of Eudontomyzon danfordi species habitat state in the studied area are: poaching, minor riverbeds morphodynamic changings, liquid and solid natural flow disruption, destruction of riparian trees and bush vegetation, habitat fragmentation-fish populations isolation, and organic/mining pollution activities.

The pattern of microhabitat preferences of Ukrainian brook lamprey Eudontomyzon mariae ammocoetes was examined in two rivers of central Poland: the Pilica River (the Vistula River basin) and the Grabia River (the Odra River basin). A comparison of abiotic factors of the rivers revealed differences in water speed and principal components: PC1 (determining gradient from decreasing medium sand to the increasing share of three fractions of gravel), PC2 (a gradient from the declining share of very coarse and coarse sand fractions to the growing content of fine sand) and PC3 (correlated with an increasing proportion of silt). The sites did not differ significantly in terms of water depth. Relative abundance and frequency of ammocoete occurrence in the Grabia River were higher than in the Pilica River. Only speed, PC1 and PC2 made a significant contribution to the prediction of larval occurrence. Eudontomyzon mariae larvae preferred substrata with a reduced amount of medium sand and increased content of gravel (PC1) as well as with a lower content of coarse sand and higher proportion of fine-grained sand (PC2). The ammocoetes also preferred areas with a water speed of 0A‚·2 m s(-1) but avoided speeds A¢Â‰Â¥ 0A‚·6 m s(-1). The abundance of E. mariae was affected by water speed, as well as by all PCs. The mean A‚± s.e. optimal current speed was 0A‚·265 A‚± 0A‚·007 m s(-1), while abundance decreased with increasing amounts of gravel (PC1) and increased with increasing amounts of fine sand and silt in the bottom substratum (PC2 and PC3). Comparison of ammocoete microhabitat use in the Pilica and Grabia Rivers showed the lack of differences in distribution in the preferred values of current speed, PC1 and PC2.

This is the first report on the fecundity in adults of two nonparasitic lamprey species at risk, the Vulnerable Kern Brook Lamprey, Lampetra hubbsi and the Critically Endangered Macedonia Brook Lamprey, Eudontomyzon hellenicus. The mean absolute fecundity of L. hubbsi was 1865 eggs and that of E. hellenicus was 994. The higher fecundity in L. hubbsi was directly related to its higher mean gonadal mass (0.90 g) relative to that in E. hellenicus (0.52 g). Based on preserved museum material possessing well-developed primary and secondary sexual characters, the spawning period for L. hubbsi was inferred to be February-March and for E. hellenicus to either extend from January to May or to occur as separate events; one in January and the other in May. Because metamorphosing ammocoetes of E. hellenicus have been collected in October and in January, it is hypothesized that those metamorphosing in October spawn in January and those metamorphosing in January spawn in May. These periods need to be corroborated with field observations. Evidence for E. hellenicus spawning in January is based on the presence of spent females of this species collected in that month.

The characteristic of distribution of Ukrainian lamprey Eudontomyzon mariae in the upper Don basin is presented. The examined part of the basin covers territories of Kursk, Orel, Tula, Ryazan, Lipetsk, Tambov, and Voronezh oblasts. It is shown that, at the present time in the upper Don basin, restoration of the range and an increase in the numbers of Ukrainian lamprey is occurring. The main concentrations of the species were recorded in the Don and Krasivaya Mecha rivers and their tributaries. In the basin of the Sosna River—one of the largest tributaries of the Don—Ukrainian lamprey is very widely distributed, but its population remains small and rarified. To preserve the species in the upper Don basin, it is suggested—in addition to already existing specially protected natural territories—to organize new ones in some small rivers and in the upper reaches of the Krasivaya Mecha River. In case of emergence of technogenic catastrophes, they can serve as donors for the entire basin of the upper Don.

C1q is the target recognition protein of the classical complement pathway and a major connecting link between innate and acquired immunity. C1q and the multifunctional tumor necrosis factor (TNF) ligand family is of similar crystal structures, are designated the C1q/TNF-related proteins (CTRPs) superfamily. They are involved in processes as diverse as host defense, inflammation, apoptosis, autoimmunity, cell differentiation, organogenesis, hibernation and insulinresistant obesity. In this study, three members of the CTRPs superfamily were isolated and characterized in Yalu River lampreys (Eudontomyzon morii), and are respectively named LaC1qC, LaCTRP1, LaCTRP9. The full-length cDNAs of C1qC-like (LaC1qAL), CTRP1-like (LaCTRP1), and CTRP9-like (LaCTRP9) consist of 723, 762 and 825 bp of nucleotide sequence encoding polypeptides of 241, 254 and 275 amino acids, respectively. All-three proteins share three common domains: a signal peptide at the N terminus, a collagenous domain (characteristic Gly-X-Y repeats), and a C-terminal globular domain. In addition, the higher expression level of the three proteins in heart by RT-PCR and real-time PCR tissue profiling implied that they might involve in immune response or injury repair of the heart in lamprey.

Fifteen new localities of the Ukrainian brook lamprey, Lampetra (Eudontomyzon) mariae, were found within 2008-2010. The majority of them are located in the Nida River drainage. Additionally, the occurrence of that species in the Strwiąz River, a tributary of the Dniester, was confirmed. The cluster analysis performed sug- gests, that L. mariae exhibits strong affinity to the stone loach, Barbatula barbatula, and the Eurasian minnow, Phoxinus phoxinus. We believe that the distribution of L. mariae on the territory of Poland seems to be strongly underestimated.

The evolution of nonparasitic brook lamprey forms and their relationship to the morphological similar parasitic river lampreys are long-standing questions in lamprey research. The present study examines the utility of 11 mitochondrial gene segments (a total of 3800 base pairs) to discriminate the closely related members of the European species pair Lamp- etra fluviatilis and Lampetra planeri. The intent was to identify fast-evolving genes in the mtDNA that might allow a dis- crimination of pair members. Therefore, the noncoding regions as the fastest evolving gene segments of animal mtDNA were also included. The results revealed that in all investigated loci, L. fluviatilis and L. planeri were genetically indistin- guishable, indicating either a very recent divergence event or ongoing gene flow between both species. The position of the Ukrainian brook lamprey, Eudontomyzon mariae, and the phylogentic relationships within the Lampetrini was inferred from partial cytochrome b by adding available sequences from GenBank to the data set. Eudontomyzon mariae formed a monophyletic cluster with Lampetra aegyptera and the European species pair. A reintegration of E. mariae in the genus Lampetra is therefore recommended. Resume´ : L'evolution des formes de lamproies de ruisseau non parasites et leur relation avec les lamproies de riviere par- asites de morphologie semblable sont des questions d'interet depuis longtemps dans la recherche sur les lamproies. Notre etude examine l'utilitede 11 segments de genes mitochondriaux (un total de 3800 paires de bases) pour distinguer les deux especes fortement apparentees de la paire de lamproies europeennes Lampetra fluviatilis et Lampetra planeri. Notre objectif est d'identifier des genes de l'ADNmt a evolution rapide qui puissent permettre de separer les membres de cette paire. Nous avons donc inclus les regions non codantes comme les segments de genes a evolution la plus rapide dans l'ADN mitochondrial chez les animaux. Nos resultats montrent que L. fluviatilis et L. planeri ne peuvent etre distingues genetiquement par aucun des locus etudies, ce qui indique soit un evenement de divergence tresr ecent ou un flux geneti- que persistant entre les deux especes. Nous avons deduit la position de la lamproie de ruisseau ukrainienne, Eudontomyzon mariae, et les relations phylogenetiques au sein des Lampetrini de l'analyse partielle du gene du cytochrome b en ajoutant des sequences disponibles dans GenBank a la banque de donnees. Euodontomyzon mariae forme un groupe monophyle ´- tique avec Lampetra aegyptera et la paire d'especes europeennes. Nous recommandons donc de reintegrer E. mariae dans le genre Lampetra. (Traduit par la Redaction)

Typically, studies of ichthyocenoses in Lithuania are being carried out to investigate the distribution, abundance and conservation status of bony fish communities. Cephalaspidomorphs as a group possess a distinctive life cycle. Therefore, researches into lampreys in Lithuania are fragmented and knowledge on species diversity, distribution, abundance and status of lamprey populations in Lithuania is poor. The present thesis reviews the distribution and abundance of Lampetra fluviatilis (Linnaeus, 1758) and Lampetra planeri (Bloch, 1784) in Lithuanian rivers, as well as the factors that affect dispersion and abundance of the larval stage of lampreys. By applying a molecular approach to identification of lamprey species, representatives of the Ukrainian lamprey Eudontomyzon mariae (Berg, 1931) were discovered inhabiting several rivers in Lithuania for the first time. A phylogenetic analysis based on mtDNA markers revealed that L. fluviatilis and L. planeri belong to the same evolutionary lineage. Additionally, the findings suggest that the Baltic Sea Lampetra spp. populations might experience the gene flow between two life strategies, which is supported by a successful in situ experimental hybridisation between L. fluviatilis and L. planeri spawning in communal redds. These findings are essential to the understanding of the population genetic structure of L. fluviatilis and L. planeri and contribute to the current knowledge of the speciation mechanism. From a practical point of view, the findings can be used in lamprey conservation. The current lamprey conservation status is assessed, and recommendations for further lamprey research and monitoring in Lithuania are given.

Typically, studies of ichthyocenoses in Lithuania are being carried out to investigate the distribution, abundance and conservation status of bony fish communities. Cephalaspidomorphs as a group possess a distinctive life cycle. Therefore, researches into lampreys in Lithuania are fragmented and knowledge on species diversity, distribution, abundance and status of lamprey populations in Lithuania is poor. The present thesis reviews the distribution and abundance of Lampetra fluviatilis (Linnaeus, 1758) and Lampetra planeri (Bloch, 1784) in Lithuanian rivers, as well as the factors that affect dispersion and abundance of the larval stage of lampreys. By applying a molecular approach to identification of lamprey species, representatives of the Ukrainian lamprey Eudontomyzon mariae (Berg, 1931) were discovered inhabiting several rivers in Lithuania for the first time. A phylogenetic analysis based on mtDNA markers revealed that L. fluviatilis and L. planeri belong to the same evolutionary lineage. Additionally, the findings suggest that the Baltic Sea Lampetra spp. populations might experience the gene flow between two life strategies, which is supported by a successful in situ experimental hybridisation between L. fluviatilis and L. planeri spawning in communal redds. These findings are essential to the understanding of the population genetic structure of L. fluviatilis and L. planeri and contribute to the current knowledge of the speciation mechanism. From a practical point of view, the findings can be used in lamprey conservation. The current lamprey conservation status is assessed, and recommendations for further lamprey research and monitoring in Lithuania are given.

Relationships among the 18 extant species of parasitic lamprey (Petromyzontiformes) were determined using a cladistic analysis of 32 mainly morphological characters. Because previous analyses support all known fossils as phylogenetically older or the same age as living lampreys, a composite agnathan fossil was used as an outgroup. A consensus of three equally parsimonious trees revealed a trichotomy between a monophyletic northern hemisphere clade and the southern hemisphere genera Geotria and Mordacia. The monophyletic status of the northern hemisphere lampreys and their classification in a single family Petromyzontidae was corroborated. It is suggested that the two southern hemisphere lamprey genera be retained as distinct families. Among northern hemisphere species, Ichthyomyzon and Petromyzon form a monophyletic group sister to the remaining genera. Caspiomyzon is sister to Tetrapleurodon + Entosphenus + Lethenteron + Eudontomyzon + Lampetra, with Tetrapleurodon in turn being sister to a grou...

The lampreys (Petromyzontiformes), one of the two surviving groups of agnathan (jawless) vertebrates, currently consist of 41 recognized species. This group has an antitropical distribution, with the 37 species of Northern Hemisphere lampreys assigned to the Petromyzontidae, whereas the four species of Southern Hemisphere lampreys are separated into either the Geotriidae (one species) or Mordaciidae (three species). All lamprey species have a blind and microphagous, burrowing larva (ammocoete), which spends a number of years in the soft sediment of creeks and rivers, after which it undergoes a radical metamorphosis. Eighteen lamprey species then embark on an adult parasitic phase (nine at sea and nine in fresh water) during which they increase markedly in size, whereas the other 23 species do not feed as adults and remain in fresh water. On the basis of morphology, 17 of the 23 non-parasitic species each evolved from a particular parasitic species whose descendants are still represented in the contemporary fauna. The remaining six non-parasitic species, the so-called “southern relict” species, have no obvious potential ancestral parasitic species, implying they have diverged markedly from their parasitic ancestor or that the parasitic ancestor is now extinct. Many of the main taxonomic characteristics reside in features that are associated with parasitic feeding, for example, the type and arrangement of the teeth on the suctorial disc and tongue-like piston. The phylogenetic relationships, derived by maximum parsimony analyses of morphological and anatomical data for the 18 parasitic species, were similar in most respects to those obtained by subjecting molecular data (cytochrome b mitochondrial DNA sequence data) for those species to Bayesian analyses. However, in contrast to the results of morphological analyses, the genera Eudontomyzon and Lampetra were not monophyletic when using molecular analyses. When non-parasitic species were included in the molecular analyses, some of the six relict non-parasitic species formed clades with parasitic species which, from their morphology, had been allocated by taxonomists to different genera. More genes, and particularly nuclear genes, should be used to help resolve the basis for these differences between the morphological and molecular phylogenies.

Eudontomyzon graecus, sp. n. (Figs. 1���2) Holotype. CMNFI 1986 ���0741: adult male, TL 136.5 mm, Filippi��s Brook, near Filippi��s, 39 �� 12 ���N 20 �� 53 ���E, Lo��ros River drainage, Ionian Sea basin, Epirus Periphery, Greece, coll. P.S. Economidis, 13 Nov. 1979. Paratypes. CMNFI 1986 ���0738: adult female, TL 146 mm, 4 ammocoetes, TL 92.5���152 mm, type locality, coll. P.S. Economidis, 27 Nov. 1969; CMNFI 1977 ��� 1741: 9 ammocoetes, TL 79���126.5 mm, type locality, coll. F. Theocharis, 17 Oct. 1973; CMNFI 1986 ���0720: 4 adult males and 6 adult females, TL 124.5��� 151 mm, 2 ammocoetes, 124���138.5 mm, type locality, coll. G. Asmis and P.S. Economidis, 20 Oct. 1977; CMNFI 1986 ���0741A: 4 adult males and 8 adult females, TL 129���158.5 mm, 4 metamorphosing ammocoetes, TL 135���156.5 mm, 3 ammocoetes, TL 151���189 mm, same collection data as holotype; ZMH 4210: 1 adult male and 1 adult female, TL 135.5���149 mm, Lo��ros River, Ionian Sea basin, Epirus Periphery, Greece, coll. P.S. Economidis, 1968. Diagnosis. Eudontomyzon graecus is distinguished from its European congeners by possessing the following character states: nonparasitic mode of life; adults with large, blunt labial teeth covering all fields in a pavement-like arrangement; one or more bicuspid teeth in the first anterial row; two or more bicuspid teeth in the first posterial row; 7���11 teeth on the infraoral lamina; endolateral formula, 2 ��� 2 ��� 2; 2���5 velar tentacles without wings; ammocoetes and adults with 53���61 trunk myomeres; rounded caudal fin; relatively longer tail length against TL; ammocoetes with relatively shorter trunk length against TL; weakly pigmented (i.e., +) upper lip; unpigmented tongue precursor bulb; unpigmented areas lateral to the tongue precursor elastic ridge. Description. Ammocoetes (Table 1). Maximum total length is 189 mm. Trunk myomeres vary from 54 to 60 with a mode of 57. Caudal fin shape is rounded in 14 individuals and spade-like in one. Lateral-line neuromasts are unpigmented and clearly appear as such by 79 mm TL. Metamorphosing ammocoetes vary in TL from 135���156.5 mm. Adults (Tables 2���3, Figs. 1���2). Total length varies from 124.5���158.5 mm. The dentition is extremely variable (Table 3). The supraoral lamina bears usually two unicuspid lateral teeth separated by a toothless bridge. In two cases, one or both lateral teeth were bicuspid. The infraoral lamina bears 7���11 teeth, usually unicuspid. In two cases, a bicuspid tooth was present. There are usually three endolateral teeth on each side of the oral disc (85 % of counts), but four teeth also occur (15 %). The most common endolateral formula is 2 ��� 2 ��� 2, but in three cases a unicuspid and in two cases a tricuspid tooth occurred. The first anterial and first posterial rows possess 4���7 teeth and 10���15 teeth, respectively, exhibiting great variability in the arrangement of the types of teeth (unicuspid, bicuspid or tricuspid), with at least one bicuspid tooth in 21 of 26 individuals in the former and 20 of 26 in the latter, in various positions in those rows. There are 3���4 rows of anterials and the same range of exolateral rows of either side of the oral disc and 1���3 rows of posterials. In eight individuals examined, including the holotype, the transverse lingual lamina exhibited an enlarged median cusp, but lateral cusps were absent and the longitudinal lingual laminae bore no discernible teeth (Fig. 2 A). Two specimens possessed what we term accessory cusps inside the ring formed by the supraoral and infraoral laminae and the left and right endolaterals. One specimen had an accessory unicuspid tooth placed at the level of the third left endolateral and the other two unicuspid teeth, one on each side below the supraoral lamina. The velar apparatus possessed 2���5 very small tentacles, with a central tentacle present in two cases and absent in three. There were no tubercles or wings. Trunk myomeres vary from 53 to 61, with strong modes of 56 and 57. In the 23 individuals, including the holotype (Fig. 1), in which the caudal fin shape could be determined, it was invariably rounded. The mean intestinal width taken at the origin of the first dorsal fin was 0.8 mm (Range: 0.5 ���1.0 mm; n = 23). Lateral line neuromasts are unpigmented. Comparative remarks. This new species belongs to Eudontomyzon Regan, 1911 because it typically possesses three lateral circumoral (= endolateral) teeth on each side of the oral disc, a prominent median cusp on the transverse lingual lamina and exolateral teeth in alate rows (Gill et al. 2003). In the adult stage, it is distinguished from E. hellenicus by having a relatively longer tail length (88 % separation based on residuals of tail length against TL; Fig. 3), usually having at least one bicuspid tooth in the first anterial row vs. none (84 % separation; Table 3) and at least two bicuspid teeth in the first posterial row vs. one or none (71 % separation; Table 3). From adults of E. stankokaramani, it is distinguished by having 7���11 teeth on the infraoral lamina vs. 5���7 (92 % separation; Table 3), a usual endolateral formula 2 ��� 2 ��� 2 vs. 1���2 ��� 1 and none of the variant formulae are shared (100 % separation; Table 3), usually 53���59 trunk myomeres vs. usually 60���65 (85 % separation; Table 3), a rounded vs. spade-like caudal-fin shape (100 % separation; this study and Holč��k & ��orić (2004), respectively) and 2���5 velar tentacles without wings vs. 9���13 (100 % separation; Table 3), of which 3���6 are folded onto the dorsal surface of the velar apparatus to form a pair of wings (Holč��k & ��orić 2004). Furthermore, the labial teeth in E. stankokaramani, when present, are very small and relatively pointed (i.e. villiform) (Karaman 1974, Holč��k & ��orić 2004) and do not fully occupy the space in the various fields (anterior, lateral and posterior), whereas in both E. graecus (Fig. 2) and E. hellenicus (Vladykov et al. 1982), they are large and blunt and form a close set pavement-like pattern that completely fills the available space in the various fields. In the ammocoete stage, it is distinguished from E. hellenicus and E. stankokaramani by having an unpigmented (i.e., ���) tongue precursor bulb vs. a usually weakly (i.e., +) pigmented bulb in E. hellenicus (91 % separation; Table 1) and a moderately to strongly (i.e., ++ and +++) pigmented bulb in E. stankokaramani (100 % separation; Table 1). Additionally, it is distinct from E. hellenicus at the ammocoete stage by having a relatively shorter trunk and longer tail (97 % separation in both cases based on residuals of trunk length against TL and tail length against TL; Figs. 4 A and 4 B, respectively), usually unpigmented areas lateral to the tongue precursor elastic ridge vs. pigmented areas (91 % separation; Table 1) and a less pigmented upper lip (vs. ��� 25 % pigment coverage giving a 91 % separation; Table 1) and from E. stankokaramani by having 54���60 trunk myomeres vs. 59���65 (95 % separation; Table 1) and a rounded vs. spade-like caudal fin (89 % separation; this study and Holč��k & ��orić (2004), respectively). While none of the characters that in combination separate E. hellenicus from E. graecus is 100 % diagnostic, the five larval characters (two morphometric and three pigmentary) that individually achieve between 91 and 97 % separation, as well as their distinct zoogeographic affinities (see below, Evolution and zoogeography), strongly suggest that they represent distinct species on different evolutionary paths. Kottelat & Freyhof (2007) stated that the new species from the Lo��ros River drainage was distinguished from Eudontomyzon hellenicus by having a longer tail (29���33 % TL vs. 26���30 % TL) and a spade-like (vs. rounded) caudal fin. The data for the tail length is taken directly from Renaud (1986) and we concur that this is correct. However, we disagree with the difference in the shape of the caudal fin. Both E. graecus and E. hellenicus almost invariably possess rounded caudal fins. In E. graecus, only one out of 15 ammocoetes (7 %) had a spade-like caudal fin and all others, as well as the 23 adults for which the shape of the caudal fin could be determined, possessed a rounded caudal fin. Likewise, 202 ammocoetes and 33 adults of E. hellenicus possessed a rounded caudal fin and only one ammocoete (E. hellenicus reaches 152 mm TL while the lamprey from the Lo��ros River basin reaches 190 mm TL. This statement is somewhat misleading and requires clarification. While it is true that the largest ammocoetes and adults of E. hellenicus are respectively 152 and 151.5 mm TL; those of the Lo��ros River drainage (E. graecus) are 189 and 158.5 mm TL (Tables 1���2). It appears that Kottelat & Freyhof (2007) were referring to the length of the adult in the case of E. hellenicus and to the length of the ammocoete in the case of E. graecus. It is significant to note that both the ammocoetes and the adults of E. graecus have a relatively longer tail length than that of E. hellenicus (Figs. 3, 4 B). This means that whatever the selective advantage conferred by the tail length, this persists after metamorphosis. Karaman (1974) and Holč��k & ��orić (2004) reported collecting metamorphosing ammocoetes of E. stankokaramani from the type locality, respectively, in February-April 1974 and 28 November 1970. The 2���5 month interval between November and February-April represents a rather long period of time for completion of metamorphosis at a single locality. According to Hardisty & Potter (1971), in the species in which metamorphosis has been studied, including the congeneric Eudontomyzon mariae (Berg, 1931), the external signs of metamorphosis are completed in about four to five weeks. A November timing of metamorphosis in E. stankokaramani is similar to that reported here for E. graecus. It is thus unfortunate that the metamorphosing individuals collected during February-April by Karaman (1974) have been lost since this represents an anomalous situation. Unfortunately, no information is available on the external pigmentation in ammocoetes and adults of E. stankokaramani because Karaman (1974) did not document it and Holč��k & ��orić (2004) stated that their specimens had probably lost their pigmentation through prolonged exposure to light. Distribution. This new species is endemic to the Epirus Periphery, Greece, being found only in the Lo��ros River and one of its tributaries, Filippi��s Brook (Fig. 5). Evolution and zoogeography. According to Economidis & Bănărescu (1991), the freshwater fish fauna of the Lo��ros River drainage and that of the Strymon River drainage belong to distinct ichthyogeographic divisions; the former to the South Adriatic-Ionian Division and the latter to the Ponto-Aegean Division, and more precisely to the Thracian-East Macedonia Subdivision. The prevalent theory for the evolution of nonparasitic species is that they are derived from a parasitic ancestor (Hubbs & Potter 1971). It is difficult, however, to speculate on the origins of the two nonparasitic species E. graecus and E. hellenicus as no parasitic ancestor has been identified through a cladistic analysis that integrates both morphological and molecular characters. A cladistic analysis based on 1,133 base pairs of the mitochondrial gene cytochrome b was recently published by Lang et al. (2009), where E. graecus (labeled as E. hellenicus from the Ionian Sea basin) and E. hellenicus constituted a clade that was sister to the Caspian lamprey, Caspiomyzon wagneri. The conclusion that the two Greek nonparasitic species are sister to C. wagneri appears premature. As mentioned in Lang et al. (2009), morphological and/or nuclear gene characters would need to be added to the cladistic analysis to further test this hypothesis. Habitat and biology. This new lamprey species spends its entire life cycle in fresh water and appears to be quite sedentary, based on the capture of all three stages of its life history (i.e., ammocoete, metamorphosing ammocoete and immature adult) at one locality in mid-November. The habitat is characterized by clear, fastflowing water and muddy substrate in which the larvae bury. The substrate may or may not include submerged vegetation. The larval stage lasts an estimated 4���6 years (Economidis 1995). Metamorphosis occurs in November based on a collection made on the 13 th of that month and would have started at least as early as 20 October based on the capture of already fully metamorphosed adults at this earlier date, but the exact duration of metamorphosis is not known. The onset of metamorphosis may be related to water temperature and summer drought. Adults do not feed as indicated by the fact that the largest ammocoetes (189 mm TL) are considerably larger than the largest adults (158.5 mm TL), coupled with the fact that the intestinal diameter of adults is 0.5 ���1.0 mm. The majority of the adult material available (24 of 26 individuals, including the holotype), collected between 20 October and 27 November, represented immature individuals. The other two adults, a male and female collected together, were sexually mature, each possessing a small (1.0 mm long) urogenital papilla. Unfortunately, their precise date of collection was not recorded, only the year. The habitat was shallow, clear water with a gravel substrate. Spawning has not been witnessed but is presumed to occur in late winter. Conservation. This new species is inherently at risk because of its very restricted distribution. Eudontomyzon hellenicus and E. graecus (then treated together as a single species) were given a status of Threatened to Endangered by Economidis (1995). A more recent evaluation of the two species (again treated as one) by Crivelli (2006) up-listed the status to Critically Endangered according to the following IUCN (2001) criteria: B 1 ab(i,ii,iii,iv,v) + 2 ab(i,ii,iii,iv,v). The situation is even more critical when one considers the two species individually. Water regulation, habitat alteration and pollution are the main threats. In the Lo��ros River drainage specifically, irrigation canals and draining ditches, often with cement-lined sides and bottom, coupled with water-level fluctuations, have resulted in reduced habitat for ammocoetes as well as reduced spawning areas for adults (Economidis 1995). Etymology. The new species name is an adjective formed from the noun Graeci, a tribe of Hellenes living since Prehistoric times in the area where the lamprey occurs. The Graeci were early on in contact with the inhabitants of the Italian Peninsula, mainly Latins, who then referred to all of the Hellenes as Greeks. Proposed Common Name: Epirus brook lamprey. A new common name is required for Eudontomyzon hellenicus as it is no longer the only Greek brook lamprey. We propose Macedonia brook lamprey as its new common name., Published as part of Renaud, Claude B. & Economidis, Panos S., 2010, Eudontomyzon graecus, a new nonparasitic lamprey species from Greece (Petromyzontiformes: Petromyzontidae), pp. 37-48 in Zootaxa 2477 on pages 38-47, DOI: 10.5281/zenodo.195327, {"references":["Regan, C. T. (1911) A synopsis of the marsipobranchs of the order Hyperoartii. The Annals and Magazine of Natural History, 7, 193 - 204.","Gill, H. S., Renaud, C. B., Chapleau, F., Mayden, R. L. & Potter, I. C. (2003) Phylogeny of living parasitic lampreys (Petromyzontiformes) based on morphological data. Copeia 2003, 687 - 703.","Holcik, J. & Soric, V. (2004) Redescription of Eudontomyzon stankokaramani (Petromyzontes, Petromyzontidae) - a little known lamprey from the Drin River drainage, Adriatic Sea basin. Folia Zoologica, 53, 399 - 410.","Karaman, M. S. (1974) Eudontomyzon vladykovi stankokaramani n. ssp. a new subspecies of lamprey from tributaries of the Ohrid-Drim-Skadar system in west Balkan Peninsula. Folia Balcanica, 3, 1 - 10, 2 figs., 6 tables.","Vladykov, V. D., Renaud, C. B., Kott, E. & Economidis, P. S. (1982) A new nonparasitic species of Holarctic lamprey, genus Eudontomyzon Regan 1911 (Petromyzontidae), from Greece. Canadian Journal of Zoology, 60, 2897 - 2915.","Kottelat, M. & Freyhof, J. (2007) Handbook of European Freshwater Fishes. Kottelat, M., Cornol, Switzerland and Freyhof, J., Berlin, Germany, 646 pp.","Renaud, C. B. (1986) Eudontomyzon hellenicus Vladykov, Renaud, Kott, and Economidis, 1982. In: Holcik, J. (Ed.), The Freshwater Fishes of Europe. AULA-Verlag GmbH, Wiesbaden, Germany, pp. 186 - 195.","Hardisty, M. W. & Potter, I. C. (1971) The general biology of adult lampreys. In: Hardisty, M. W. & Potter, I. C. (Eds.), The Biology of Lampreys. Vol. 1. Academic Press, London, U. K., pp. 127 - 206.","Berg, L. S. (1931) A review of the lampreys of the northern hemisphere. Annuaire du Musee Zoologique de l'Academie des Sciences de l'URSS, 32, 87 - 116 + 8 pls.","Economidis, P. S. & Banarescu, P. M. (1991) The distribution and origins of freshwater fishes in the Balkan Peninsula, especially in Greece. Internationale Revue der gesamten Hydrobiologie und Hydrographie 76, 257 - 284.","Hubbs, C. L. & Potter, I. C. (1971) Distribution, phylogeny and taxonomy. In: Hardisty, M. W. & Potter, I. C. (Eds.), The Biology of Lampreys. Vol. 1. Academic Press, London, U. K., pp. 1 - 65.","Lang, N. J., Roe, K. J., Renaud, C. B., Gill, H. S., Potter, I. C., Freyhof, J., Naseka, A. M., Cochran, P., Espinosa Perez, H., Habit, E. M., Kuhajda, B. R., Neely, D. A., Reshetnikov, Y. S., Salnikov, V. B., Stoumboudi, M. Th. & Mayden, R. L. (2009) Novel relationships among lampreys (Petromyzontiformes) revealed by a taxonomically comprehensive molecular data set. In: Brown, L. R., Chase, S. D., Mesa, M. G., Beamish, R. J. & Moyle, P. B. (Eds.), Biology, Management, and Conservation of Lampreys in North America. American Fisheries Society, Bethesda, Maryland, U. S. A., pp. 41 - 55.","Economidis, P. S. (1995) Endangered freshwater fishes of Greece. Biological Conservation, 72, 201 - 211.","Crivelli, A. J. (2006) Eudontomyzon hellenicus. In: IUCN 2010. IUCN Red List of Threatened Species. Version 2010.1. Available from: http: // www. iucnredlist. org / (31 March 2010)."]}

Key to larvae and adults of Greek brook lampreys of the genus Eudontomyzon 1 Horseshoe-shaped oral hood with oral cirri inside (larva)........................................................................................... 2 ��� Oral disc with fimbriae and papillae at its periphery and teeth inside (adult)............................................................. 3 2 Upper lip pigmentation .................................................................. Eudontomyzon graecus ��� Upper lip pigmentation ��� 25 % coverage; tongue precursor bulb and both sides of elastic ridge pigmented; trunk length 48.2���55.4 % TL; tail length 21.2���30.6 % TL .............................................................. Eudontomyzon hellenicus 3 First anterial row with one or more bicuspid teeth; first posterial row with two or more bicuspid teeth; tail length 29.3 ���33.0% TL ......................................................................................................................... Eudontomyzon graecus ��� First anterial row teeth all unicuspid; first posterial row teeth all unicuspid or with a single bicuspid tooth; tail length 26.2���30.4 % TL ..................................................................................................................... Eudontomyzon hellenicus, Published as part of Renaud, Claude B. & Economidis, Panos S., 2010, Eudontomyzon graecus, a new nonparasitic lamprey species from Greece (Petromyzontiformes: Petromyzontidae), pp. 37-48 in Zootaxa 2477 on page 38, DOI: 10.5281/zenodo.195327

After World War II two fish species (Acipenser sturio and Salmo salar) became extinct in Poland. At present there are 8 most endangered species, mostly anadromous and ecologically highly specialized (Petromyzon marinus, Eudontomyzon mariae, Lampetra fluviatilis, Salmo trutta lacustris, Pelecus cultratus, Moroco percnurus, Gobio kessleri, and G. albipinnatus). As much as 58.9% of fishes and lampreys are threatened to various extents. Conservation of species that are going extinct depends upon artificial spawning and stocking, with simultaneous improvement of water quality and prevention of destruction of aquatic habitats.

Eudontomyzon lanceolata (Kux & Steiner, 1972) Inland water: 31400-806 (1 spa), 01.07.1992, Of Stream, Rize-Trabzon; 31400-659 (1 spa), 02.04.2004, Iyi Stream, Rize., Published as part of Nurettin Meri��, L��tfiye Eryilmaz & M��fit ��zulug, 2007, A catalogue of the fishes held in the Istanbul University, Science Faculty, Hydrobiology Museum., pp. 29-54 in Zootaxa 1472 on page 31

Eudontomyzon lanceolata (Kux & Steiner, 1972) Inland water: 31400-806 (1 spa), 01.07.1992, Of Stream, Rize-Trabzon; 31400-659 (1 spa), 02.04.2004, Iyi Stream, Rize.

The Ukrainian brook lamprey Eudontomyzon mariae was collected in two streams belonging to the Sava river basin in Croatia. The great variation in morphological characters of this species is emphasized and its description is augmented. Clinal variation in the trunk myomeres, total length and the oral disc dentition is detected.

Eudontomyzon graecus, sp. n., a nonparasitic lamprey from Greece, is restricted to the Loúros River drainage, Ionian Sea basin, Epirus Periphery. It is distinguished from Eudontomyzon hellenicus, the other nonparasitic Greek brook lamprey, occurring allopatrically in the Strymon River drainage, Aegean Sea basin, Central Macedonia Periphery, by possessing at the adult stage a relatively longer tail and a greater number of bicuspid teeth in both the first row of anterials and posterials, and at the ammocoete stage, by a relatively shorter trunk and longer tail and a less pigmented upper lip, tongue precursor bulb and lateral aspects of the elastic ridge. Taken individually, these three adult and five larval characters, varied in their diagnostic power from 71 to 88% for the adults and from 91 to 97% for the ammocoetes. However, in combination, these characters provide sufficient evidence to justify recognition of this new species, in addition to the wide distance (ca 400 km) separating it from its closest extant relative, as well as its independent origin.

The distribution of living lampreys (Petromyzoniformes) is antitropical, a feature almost certainly related to the fact that the ultimate incipient lethal temperature of ammocoetes lies between ~28 and 32 °C. In general, the species that attain the largest body size have the widest distribution. The holarctic species are placed in the Petromyzonidae, while those of the Southern Hemisphere are assigned either to the Geotriidae or Mordaciidae. Reference is made to the techniques used in lamprey systematics, placing particular emphasis on defining the terminology of the components of the dentition and the ways in which body proportions can be measured. Using dentitional characters, the Petromyzonidae can be arranged in a logical sequence from Ichthyomyzon and Petromyzon to either Caspiomyzon or to series consisting either of Eudontomyzon and Tetrapleurodon or the subgenera of Lampeira, i.e. Entosphenus, Lethenteron, and Lampetra. A list is given of the 17 parasitic and 22 nonparasitic lampreys described from the Northern and Southern hemispheres. The evolution of nonparasitic lampreys has involved an extension of larval life and a reduction in the postmetamorphic period, a process which has led to a marked reduction in the size of the mature adults. The more conspicuous differences between nonparasitic and parasitic species start to appear after the onset of metamorphosis. These include the rapid enlargement and maturation of gonads in nonparasitic species. However, significant differences have been found between the larvae of nonparasitic lampreys and those of their parasitic ancestor in such features as the number of oocytes, trunk myomeres, and size at the onset of metamorphosis.Key words: lampreys, life cycles, parasitic, nonparasitic, distribution, dentition, phylogeny, Petromyzonidae, Geotriidae, Mordaciidae

A new nonparasitic lamprey, Eudontomyzon hellenicus, from the Struma and Loúros river basins in Greece, is described and illustrated. The holotype (No. NMC 77-1773) is deposited in the National Museum of Natural Sciences, Ottawa, Canada. From all species of the same genus, E. hellenicus differs in the following characters: (1) small metamorphosed size (maximum total length (TL) 159 mm), (2) fewer trunk myomeres (52–62, averaging 57.4), (3) poorly developed lingual laminae cusps, (4) short and few (2–5) velar tentacles, and (5) extremely short urogenital papilla, even in spawning males. Furthermore, a nonfunctional intestinal tract in metamorphosed individuals of our species separates it from parasitic E. danfordi. The description is based on a study of 64 metamorphosed specimens, 13 transforming individuals and 237 ammocoetes.

The new term 'satellite species' is used to replace the less accurate and by now obsolete term 'paired species.' Among lamprey genera a complex of species is usually found: one parasitic or 'stem' species and one or several nonparasitic or 'satellite' species probably derived from it. Among eight genera of holarctic lampreys the interrelation between the number of parasitic (p) and satellite (s) species is as follows: Caspiomyzon, 1p and no s; Petromyzon, 1p and no s; Ichthyomyzon, 3p and 3s; Tetrapleurodon, 1p and 1s; Entosphenus, 3p and 3s; Lampetra in Europe, 1p and 2s, in North America, 1p and 3s; Eudontomyzon in Europe, 1p and 3s, in Asia, 1p and no s; Lethenteron, 1p and 5s. Thus among the six polyspecific genera of the holarctic lampreys, 12 parasitic and 20 satellite species are presently known.

Summary The Autor refers about the first results of his own research on Lampreys of Austrian Scientificic Institutes and Museums, particularly on the Lampreys of the Danube basin. He points out that the Lampreys wich he found classified as L. fluviatiis and L. planeri, are instead to be considered as Eudontomyzon danfordi (Regan). Only two specimens, fished in the Inn river (Higher Austria), meight eventually belong to L. planeri. Thus it results that E. danfordi lives in the main and lower cours of the Danube (from Linz to the mouth), in the Raab and Drava rivers, whereas up to the present time specimes of this species fished in the lower course of the Danube ware the only ones to be known. Finally the Author speaks of this Lamprey's biology.

Evidence is presented which suggests that the nonparasitic lamprey. Lampetra (Eudontomyzon) gracilis Kux, 1965, is conspecific with the parasitic lamprey Eudontomyzon danfordi Regan, 1911. The diagnostic characters of the holotype and of the non-type material of E. gracilis arc features found in E. danfordi specimens in their second and final year of adult life, thereby making the former a junior synonym of the latter.

The Carpathian brook lamprey (Eudontomyzon danfordi Regan, 1911) is an endemic protected species of Cephalaspidomorphi in the Carpathian Basin. No parasites have become known from these jawless vertebrates to date. Here, the authors describe an infection from a single specimen manifesting in protuberant skin cysts 7-10 mm in diameter, scattered on the body surface. Similar dermal infection was observed in 25 of the 274 lampreys recorded in the population survey. Skin cysts filled with round spore-like structures of a dermocystid parasite were found. These particles measured 8-14 µm in diameter and had an about 0.5 µm thick wall, and containing mainly a granular mass and a relatively scarce plasma. No hyphae were recorded. Despite conspicuous morphological changes in the skin, no inflammatory reactions were found. The molecular analysis of 18S rDNA showed similarity to dermocystid species of several fish species but differed from them approximately by 2%. This is the first record of a dermocystid parasite infecting a jawless vertebrate.

The larval lamprey (ammocoete), a relatively sedentary animal living in burrows in the soft deposits of streams and rivers, relies for food on the filtration of microscopic detritus and plant and animal organisms. The water current entering the pharyngeal chamber serves a nutritive as well as a respiratory function, the food particles being collected on mucous strands which are passed backwards to the intestine (Sterba 1953, 1961). While some workers have recorded the presence of Protozoa, rotifers, nematodes and even small annelids in the gut (see Hardisty & Potter 1971), there can be little doubt that algae, and particularly diatoms, are much more frequently ingested (Creaser & Hann 1929; Schroll 1957, 1959; Alvarez del Villar 1966; Moore 1972; Potter, Cannon & Moore in press). That algae are of nutritional value is demonstrated by the fact that the proportion of cells with colourless chloroplasts is significantly higher in the posterior region of the gut (Moore & Beamish 1973). In this latter investigation of the feeding biology of the larvae of the landlocked and anadromous forms of the sea lamprey, Petromyzon marinus L., a relationship was shown to exist between the number of algal cells in the gut and both the size of the animal and the temperature of the water. It also demonstrated that, particularly in the winter, a considerable amount of detritus was ingested, an observation also made by Schroll (1959) in the case of the brook lampreys Lampetraplaneri (Bloch) and Eudontomyzon danfordi Regan. Detritus, it should be noted, is regarded in this study as finely particulate decaying plant and animal material. Although there is no information on the digestive efficiency of the ammocoete, the presence of large numbers of live algal cells in the hind end of the gut suggests that it is almost certainly low. Furthermore, the possibility cannot be excluded that, in addition to algae and detritus, bacteria and the organic substances dissolved in the water are also of significance as a food source. The view that the latter two may be relevant is supported by the observation that relatively high densities of ammocoetes have been reported in places where there are decaying animal carcasses (Hardisty & Potter 1971). This paper reports experiments carried out in the laboratory to ascertain the extent to which ammocoetes of the brook lamprey, Lampetraplaneri, can utilize algae, detritus, bacteria and dissolved organic substances for growth. Determinations have also been made of the assimilation efficiency of larvae fed exclusively on a diet of algae.