Revue suisse de Zoologie (March 2016) 123(1): 1-9 ISSN 0035-418 An emendation of the generic diagnosis of the monotypic Glanitaenia (Cestoda: Proteocephalidae), with notes on the geographical distribution of G. osculata, a parasite of invasive weis catfish Alain de Chambrier1 & Tomäs Scholz2-* ' Département des Invertébrés, Muséum d'histoire naturelle, CP 6434, CH-1211 Genève 6, Switzerland. E-mail: alain. dechambrier@ville-ge. ch 2 Institute of Parasitology, Biology Centre of the Czech Academy of Sciences, Branisovskd 31, 370 05 Ceské Budëjovice, Czech Republic. E-mail: [email protected] Corresponding author. E-mail: [email protected] Abstract: The generic diagnosis of the monotypic Glanitaenia is amended based on a detailed morphological examination of newly collected specimens of G. osculata (Goeze, 1782) (syn. Proteocephalus osculatus) found in weis catfish, Silurus glanis Linnaeus, 1758, from Neuchâtel, Switzerland. Several morphological characteristics of G. osculata are described for the first time or better specified, such as uterine development (type 2 according to the classification by de Chambrier et al, 2004, 2015), the presence and peculiar position of a vaginal sphincter (previously not reported), its egg morphology, description of scolex microtriches and a dense network of osmoregulatory canals in the posterior part of the scolex and the anterior region of the neck (proliferative zone). The anterior position of the vagina, which opens anterior to the cirrus-sac, not ventral as typical for most of the closely related species of the Proteocephalus aggregate, may represent apomorphy of this taxon. Glanitaenia osculata is reported from Italy (River Po basin), Romania (River Danube delta) and Switzerland (Aare/Rhine River basin) for the first time. This expansion of the parasite distribution area may be related to a recent introduction of weis catfish to West Europe. Keywords: Cestoda, Glanitaenia osculata, morphology, redescription, freshwater fish, geographical distribution, Europe. INTRODUCTION Glanitaenia was differentiated from the other genera of the subfamily Proteocephalinae by the possession of The cestode genus Glanitaenia de Chambrier, a well-developed, functional apical sucker with a deep Zehnder, Vaucher & Mariaux, 2004 (Proteocephalidea: cavity (the apical sucker in other species of the Proteo- Proteocephalidae) was erected by de Chambrier et al. cephalus-aggregate is vestigial, without any cavity, or (2004) based on the phylogenetic position of its type completely absent - see Scholz et al, 1998). In the dia¬ and only species, G. osculata (Goeze, 1782) [syn. gnosis of Glanitaenia, de Chambrier et al (2004) relied Taenia osculata Goeze, 1782; Proteocephalus osculatus mainly on the morphological features of G. osculata de¬ (Goeze, 1782) La Rue, 1911; Gangesia osculata (Goeze, scribed by previous authors, especially Nybelin (1942), 1782) Reichenbach-Klinke, 1962] from weis catfish, Freze (1965), Scholz & Hanzelovâ (1998), and Scholz et Silurus glanis Linnaeus, 1758, among Palaearctic al. (1998). However, some morphological details, includ¬ proteocephalideans. It grouped with morphologically ing characteristics that are recently considered to be of distinct Paraproteocephalus par asiluri (Yamaguti, potential taxonomic and phylogenetic importance, such 1934), another parasite of si lurid catfishes, thus as the development of the uterus, relative size of the ova¬ making the genus Proteocephalus sensu stricto (called ry, morphology of the eggs and terminal genitalia (the Proteocephalus-aggregate by de Chambrier et al, 2004) distal part of the vaginal canal), and the course of osmo¬ paraphyletic. The erection of Glanitaenia as a separate regulatory canals (see de Chambrier et al, 2012, 2015), genus was supported by phylogenetic analyses based on were not provided. 5.8S + ITS2, V4 region of the 18S rRNA and 28S rRNA Therefore, the generic diagnosis of Glanitaenia is gene sequences (Hypsa et al, 2005; Scholz et al, 2007; amended in the present paper and the current distribution de Chambrier et al., 2015). of G. osculata is reviewed based on new geographical Manuscript accepted 06.07.2014 DOI: 10.528 l/zenodo.46282 2 A. de Chambrier& T. Scholz records from Europe and in relation to the current (ii) tapeworms from the Orlik water reservoir on the River expansion of its fish host, weis catfish, which is classified Vltava at Stedronin, South Bohemia, Czech Republic, as an invasive species (Copp et al, 2009). collected on 12 May 1996 (8 specimens; host field No. FM5504), 18 September 1996 (1 spec.), 22 May 1997 (6 spec.; Nos. 2146 & 2147), 4 June 1997 (34 spec.; No. MATERIAL AND METHODS 2186), 11 June 1997 (2 spec.; Nos. 2241-2243) and 13 October 1997 (8 spec.; Nos. 2244 & 2245) (see Scholz et The present study is mainly based on morphological eval¬ al, 1998; Scholz & Hanzelovâ, 1998); uation of 25 tapeworms found in a single weis catfish, (iii) 1 specimen from Horusickÿ fishpond. South Silurus giants Linnaeus, 1758 (total length of 211 cm, Bohemia, Czech Republic (Moldau/Elbe River basin) weight of 68 kg) from Neuchâtel, Switzerland (River collected on 19 October 2009; Aare basin) examined on 25 March 2010 by the senior (iv) 4 specimens from Bodensee, Germany, collected in author (A. de C.). Specimens for morphological study 1999 by Alexander Brinker; were fixed immediately after dissection of the host with (v) 1 specimen from River Po in Ferrara, Italy, collected hot 4% neutral formaldehyde solution and then stored in July 2009 by Bahram S. Dezftili; in 75% ethanol. They were stained with Mayer’s hydro¬ (vi) 1 specimen from Chilia Branch of the Danube River, chloric cannine solution, dehydrated in a graded ethanol Romania, collected in July 2013 by Petr Horâk (all series, cleared with eugenol (clove oil) and mounted in specimens listed above - IPCAS C-49/1). Canada balsam. Pieces of strabila were embedded in pa¬ Relative size of the ovary, i.e. the proportion of its size raffin wax, transversely sectioned at 12-15 pm, stained in relation to the size of the proglottid, was calculated with Weigert’s haematoxylin and counterstained with 1% as outlined by de Chambrier et al. (2012). A copy of the eosin B (acidified with five drops of pure acetic acid for illustration of the proglottid was cut out and weighed 100 ml solution) (de Chambrier, 2001; Oros et al, 2010). using a high precision balance (Mettler AE 163). Then Voucher specimens have been deposited in the collec¬ a copy of the ovary itself was cut out and weighed, and tion of the Natural History Museum in Geneva (acro¬ the proportion of its weight to that of the entire proglottis nym MHNG-PLAT; Collection numbers 68395-68397, was calculated (in %). 68699, 68700, 68709, 84707-84712, 91260) and in the helminthological collection of the Institute of Parasitolo¬ gy, Biology Centre of the Czech Academy of Sciences in RESULTS Ceské Budëjovice (acronym IPCAS; Collection number C-49/1). Examination of newly collected cestodes from Scoleces of two specimens were prepared for scanning Switzerland as well as voucher specimens from the electron microscopy (SEM) as follows: specimens were Czech Republic, Germany, Hungary, Italy, Romania dehydrated in a graded ethanol series, then transferred and Slovakia made it possible to supplement the to a graded amylacetate series, critical point-dried in morphological description of Glanitaenia osculata, with CO„ sputter-coated with gold and examined in a Zeiss morphological data, which were previously not reported DSM 940A electron microscope at the Natural History or not described sufficiently (Figs 1-15). Museum, Geneva. Microthrix terminology follows that of Chervy (2009). 1. Development of the uterus Eggs were studied in the water after their spontaneous The uterine development of G. osculata is of type 2 release from gravid tapeworms found in S. glanis from the according to the classification of de Chambrier et al. River Vltava at the Orlik water reservoir near Stedronin, (2004, 2015). In immature proglottids, the uterine stem Czech Republic, examined by T. S. on 13 October 1997 is lined by numerous weakly-staining cells, sometimes (host field nos. 2244 & 2245 - see below) and used difficult to observe, on both sides. Thereafter, elongated for experimental infection of copepods (see Scholz, ramified digitations appear lateral to the stem; their 1999). Measurements of egg envelopes, oncosphere and apical part contains undifferentiated chromophilic cells. embryonic hooks are expressed in micrometres (pm) as In the last immature proglottids, the lumen of the uterine range with the mean ± standard deviation and number of stem occupies up to 34% of the proglottid width. In measurements in parentheses. mature proglottids, the lumen gradually extends from In addition, the following comparative material of its base to the apex of each digitate diverticulum and G. osculata from S. glanis deposited in IPCAS was occupies up to 44% of the proglottid width; the apex studied: is lined with numerous chromophilic cells. Lateral (i) 5 specimens from the Rivers Latorica in Slovakia and ramified diverticula enlarge in gravid proglottids, Tisa near Szolnok and Kötelek (Hungary) (see Scholz, occupying up to 59% of the proglottid width (Figs 10, 1989); 11). Amended diagnosis of G/anitaenia 3 wmm Figs 1-9. Glanitaenia osculata from Silurus glanis, Switzerland. 1-7. Scanning electron micrographs (INVE-PLAT-91260). (1) Scolex, dorsoventral view. (2) Scolex, apical view. (3) Scolex, lateral view. (4) Acicular filitriches on the apex of the scolex. (5) Papilliform filitriches on the internal surface of suckers. (6) Capilliform filitriches between the suckers. (7) Gladiate spinitriches on the external surface of the proliferation zone (neck). (8, 9.) Photomicrographs of longitudinal sections of the scolex (INVE-PLAT-91260). (8) Detail of the apical part of the scolex showing an apical sucker. (9) Detail of the dense network of osmoregulatory canals in the posterior part of the scolex and the anterior part of the neck (proliferation zone). 4 A. de Chambrier& T. Scholz tg St SU cc ud im vo vi Figs 10, 11. Glanitaenia osculata from Silurus glanis, Switzerland. (10) Pregravid proglottid, ventral view (MHNG-PLAT-84710). Note the shape of the ramified lateral diverticula of the uterus. (11) Cross section of a pregravid proglottid at the level of the preporal region. Abbreviations: cc: chromophilic cells; do: dorsal osmoregulatory canal; 1m: internal longitudinal musculature; mg: Mehlis’ glands; ov: ovary; st: subtegumental muscle fibres; su: subtegumental cells; te: testes; tg: tegument; ud: uterine diverticula; us: uterine stem; vi: vitelline follicles; vo: ventral osmoregulatory canal. Amended diagnosis of G/anitaenia 5 mf vs 250 [irn 250 pm 25 [jm mf VS 500 pim Figs 12-15. Glanitaenia osculata from Silurus glanis, (12, 13) Cross sections of the poral part of a pregravid proglottid at the level of the vagina (MHNG-PLAT-68397). Note the presence of a proximally situated vaginal sphincter and more distally situated muscles surrounding the vaginal canal. (14) Egg spontaneously released from a gravid tapeworm collected in the Czech Republic. (15) Detail of the cirrus-sac and the distal (terminal) part of the vaginal canal (MHNG-PLAT-84710). Abbreviations: ci: cirrus; cs: cirrus-sac; em: bi-layered embryophore; ga: genital atrium; 1m: internal longitudinal musculature; mf: muscle fibres; mi: microtriches; oe: outer envelope; on: oncosphere; st: subtegumental muscle fibres; su: subtegumental cells; te: testes; tg: tegument; va: vas deferens; vc: vaginal canal; vi: vitelline follicles; vo: ventral osmoregulatory canal; vs: vaginal sphincter. 6 A. de Chambrier& T. Scholz 2. Relative size of the ovary 8, Microtriches The surface of the ovary of G. osculata represents The apical part of the scolex is covered with acicular 7.5-8.7% of the total surface of the proglottids in the filitriehes (Fig. 4) and the internal surface of the newly collected material, and 10.6-10.7% in proglottids suckers with papilliform filitriehes (Fig. 5); capilliform illustrated by Scholz & Hanzelovâ (1998) (calculated filitriehes are present between the suckers (Fig. 6) from fig. 17A, C in that paper). and gladiate spinitriches on the external surface of the proliferation zone (neck) (Fig. 7). 3. Vaginal sphincter On the basis of our new observations, we amend the diagnosis of Glanitaenia as follows: The distal part of the vaginal canal of G. osculata is surrounded by a small circular sphincter. Unlike most Glanitaenia de Chambrîer, Zehnder, Vaucher & other proteocephaline cestodes, it is situated more Mariaux, 2004 - amended diagnosis medially, ventral or medioventral to vitelline follicles (Figs 12, 13, 15), not close to the genital atrium as in Diagnosis: Proteocephalidea, Proteocephalidae. Lar¬ other taxa. In addition to this circular sphincter of a ge tapeworms with numerous slightly craspedote typical appearance (ring-like, circular sphincter), the proglottids, wider than long including for pregravid and distal (terminal) part of the vaginal canal is surrounded gravid proglottids. Inner longitudinal musculature well by a few separated bundles of muscle fibres (Figs 12, developed, formed by highly anastomosed, numerous 13, 15). bundles of muscle fibres. Scolex unarmed, with four uniioculate suckers, with convoluted osmoregulatory canals in its posterior part. Apical sucker well developed, 4. Position of the vaginal canai strongly muscular, with deep cavity; apical part lined with The vaginal canal is almost always anterior to the numerous gland cells. Neck region (proliferation zone) cirrus-sac (in 163 of 166, i.e. 98.1%, of observed long, containing extensively developed, convoluted proglottids; Fig. 15), thus not overlapping it on its osmoregulatory canals with numerous anastomoses in ventral side. In most species of the Proteocephalus its anterior part. Immature proglottids numerous, begin aggregate, the vaginal canal opens at the same level as at considerable distance posterior to scolex. Testes the cirrus pore, thus the distal part of the vaginal canal numerous, medullary, in one continuous field and in overlaps the terminal part of the cirrus-sac on its ventral one layer, exceptionally with a few testes in second side (Scholz & Hanzelovâ, 1998). incomplete layer. Ovary medullary, numerous, bilobed, relatively large in relation to surface of proglottids. Vagina opening to common genital atrium anterior to cirrus-sac; 5. Egg morphology vaginal canal with circular vaginal sphincter situated ventral to ventrolateral to vitelline follicles; terminal Eggs released spontaneously to the water are spherical, (distal) part of vaginal canal encircled by separated with a hyaline outer envelope, 34-43 (39.3 ± 2.3; n = bundles of muscle fibres. Genital pore slightly pre- 27) in diameter; diameter of granular embryophore equatorial. Vitelline follicles lateral, occupying almost 23-27 (25.3 ± 1.2; n = 24) (Fig. 14). Oncosphere entire proglottid length. Uterus medullary, ventral, with subspherical, 15-18 (16.7 ± 0.9; n = 20) long and 14-17 ramified lateral diverticula. Development of uterus of (14.7 ± 0.8; n = 20) wide, with three pairs of embryonic type 2. Parasites of siluriformi fish in Palaearctic Region. hooks (Fig. 14); lateral hooks slightly shorter, 6.5-7.7 Type and only species: Glanitaenia osculata (Goeze, (7.1 ± 0.3; n = 25) long, than median hooks, 6-7.5 (6.7 ± 1782) de Chambrier, Zehnder, Vaucher & Mariaux, 2004 0.4; n = 20) long. [synonyms Taenia osculata Goeze, 17'82; Proteocephalus osculatus (Goeze, 1782) La Rue, 1911; Ichthyotaenia skorikowi von Linstow, 1904; Proteocephalus skorikowi 6. Osmoregulatory canals (von Linstow, 1904) La Rue, 1911; Gangesia osculata In the posterior part of the scolex and the anterior part (Goeze, 1782) Reichenbach-Klinke, 1962]. of the neck region of G. osculata, osmoregulatory canals are strongly convoluted and form a dense network of anastomosed canals (Fig. 9). DISCUSSION In the present study, generic diagnosis of one of the earliest diverging taxon of the subfamily 7. Number of testes Proteocephalinae (see de Chambrier et al, 2015) is The number of testes in the newly collected specimens amended, based on the examination of newly collected is 222-281 per proglottid (x = 249; n = 10; Fig. 10). and museum material of its type and only species, Amended diagnosis of Glanitaenia 1 Glanitaenia osculata. This species was poorly known section at the level of the vagina), but these cells were not since its description as Taenia osculata by Goeze (1782) interpreted as a vaginal sphincter. Similarly, no sphincter and La Rue (1911, 1914) placed it (as Proteocephalus was illustrated in the terminal part of the vaginal canal osculatus) among species inquirendae, partly because (fig. 161 in that paper). it was confused with another specific parasite of One of the reasons why Scholz & Hanzelovâ (1998) did weis catfish, Silurotaenia siluri (Batsch, 1786) not report the vaginal sphincter, which is actually present (Proteocephalidea: Gangesiinae). Nybelin (1942) was in the specimens studied by these authors, could be its the first who confirmed the validity of P. osculatus and unusual position. It is situated at the level of vitelline provided a most detailed description of its morphology follicles, i.e. much more medially than in related taxa at a given time. such as Proteocephalus longicollis (Zeder, 1800) and Freze (1965), who studied extensive material of P percae (Müller, 1780), in which the sphincter is close P osculatus (= Glanitaenia osculata) from the former to the genital atrium (see figs 14G, 18E, F and 20 H, I in Soviet Union, stated that “Nybelin’s description of Scholz & Hanzelovâ, 1998). P. osculatus [= G. osculata] insufficiently reflects the The terminal part of the vaginal canal of G. osculata, boundaries of its morphological variations, although which is surrounded by a medially situated circular Freze’s material in general corresponds to this description vaginal sphincter of a typical shape (ring-like sphincter) and pertains to the same species.” Scholz et al. (1998) and a few separated bundles of muscle fibres situated more provided detailed data on the scolex morphology of distally, somewhat resembles that of phylogenetically G. osculata (as P. osculatus) including SEM micrographs, distant species from Neotropical catfish, Mariauxiella and Scholz & Hanzelovâ (1998) redescribed the species piscatorum de Chambrier & Vaucher, 1999 (see fig. 87 in based on newly collected specimens from the Czech de Chambrier & Vaucher, 1999). Republic. The eggs of G. osculata are described and illustrated for In the present study, some morphological and biometrical the first time in the present paper, even though Scholz data are provided for the first time, which made it (1999) studied early phases of the developmental possible to amend the generic diagnosis of Glanitaenia. cycle of the species. However, he provided only data de Chambrier et al. (2004) have demonstrated potential on metacestodes (plerocercoids) from experimentally importance of the uterine development as one of very infected copepods 12 and 21 days post infection. In fact, few morphological characteristics that may reflect the eggs of G. osculata resemble those typical of most the evolutionary history of the order (see also de species of the Proteocephalus aggregate (see Scholz, Chambrier et al., 2015). The former authors classified the 1999). development of the uterus of G. osculata as type 2 (see The anterior part of the body of G. osculata, specifically fig. 1 in de Chambrier et al., 2004), but did not provide the posterior part of its scolex and the anterior part of any details. Surprisingly, the shape of the developed a very long proliferative zone (neck region), contains uterus of G. osculata with ramified lateral diverticula is strongly convoluted osmoregulatory canals that form most similar to that of Ageneiella brevifilis de Chambrier a dense network of anastomosed canals. Scholz et al. & Vaucher, 1999, a parasite of the auchenipterid catfish (1998) observed similar networks in the scolex of most Ageneiosus inermis (Linnaeus) in the Neotropical Region Palaearctic species of the Proteocephalus aggregate, but (see fig. 31 in de Chambrier & Vaucher, 1999), which they did not focus on the presence of these canals in the belongs to a most derived clade of proteocephalidean proliferative zone. In addition, the canals seem to be cestodes (de Chambrier et ai, 2015). much more developed in G. osculata compared to those The new material of G. osculata also enabled us to in species of the Proteocephalus aggregate (see figs 1K, provide data on the relative size of the ovary (see de 3G, I, J, 5E, Q in Scholz et ai, 1998). A dense network Chambrier et ai, 2012). The surface of the ovary of of osmoregulatory canals situated in the posterior part of G. osculata represents 7.5-8.7% of the total surface of the scolex and the anterior part of the neck region was proglottids (and 10.6-10.7% in specimens illustrated by Scholz & Hanzelovâ, 1998), which well corresponds to also observed in phylogenetically distant Proteocephalus the values in other species of proteocephalideans from regoi de Chambrier, Scholz & Vaucher, 1996, a parasite teleost fishes and it is markedly larger than those in most of Hoplias malabaricus (Bloch) (Characiformes: species of Ophiotaenia La Rue, 1911 from reptiles (see Erythrinidae) in the Neotropical Region (see figs 1 and 3 table 1 in de Chambrier et ai, 2015). in de Chambrier et ai, 1996), and in Sandonella sandoni A novelty of the present study is a detailed description of (Lynsdale, 1960) from Heterotis niloticus (Cuvier) a vaginal sphincter in G. osculata and its peculiar position (Osteoglossiformes: Arapaimidae) in the Ethiopian and morphology. Indeed, Nybelin (1942) also described Region (see fig. 2 in de Chambrier et ai, 2008). Function a small vaginal sphincter and illustrated it in his fig. 3, of these canals and the reason of their concentration in but he did not provide any details. In contrast, Scholz the scolex or in the proliferative zone are not known. & Hanzelovâ (1998) did not report this structure at all. Cestodes seem to be osmoconformers whose excretory They illustrated a thick layer of cells lining the terminal system plays little or no role in osmoregulation and is (distal) part of the vaginal canal in their fig. 9G (cross largely excretory (Smyth & McManus, 1989). 8 A. de Chambrier& T. Scholz Glanitaenìa osculata belongs to the largest cestodes in G. osculata versus L-shaped, with most follicles of teleost fishes, with a total length reaching up to 1 m forming a transverse band lateral to the ovary in the latter (Lühe, 1910); the longest specimen found in the present species - see figs 6 & 7 in Shimazu, 1993), and uterine study was 74 cm long (a tapeworm from Switzerland). diverticula (lateral in the former species as in a majority The body of G. osculata consists of numerous, rather of proteocephalideans versus forming anterior and large proglottids that contain many testes. Even though posterior branches from the transversely situated uterine these are mostly in a single layer, their counting is stem in P. parasiluri -figs 6 & 7 in Shimazu, 1993). difficult due to their high number and a well-developed Glanitaenia osculata resembles species of the longitudinal musculature. Nybelin (1942) reported 276- Proteocephalus aggregate in the morphology of its 291 testes per proglottid, Freze (1965) 180-424 testes scolex, which is devoid of any metascolex, and shape of (usually about 250), but Scholz & Hanzelovâ (1998) the strabila and its proglottids, which are only slightly only 105-171 testes. In the present study, 222-281 testes wider than long. It can be distinguished from species of (x = 249, n = 10) were counted based on illustrations of the Proteocephalus aggregate by a much more developed individual proglottids, which is the most precise method apical sucker (versus vestigial or absent in species of the of counting the testes. latter group - see Scholz et al., 1998) and by the anterior This species forms, together with Paraproteocephalus position of the vaginal canal in relation to the cirrus-sac parasiluri (Yamaguti, 1934), type and single species (ventral to the terminal part of the sac in most species of of Paraproteocephalus Chen in Dubinina, 1962 (see the Proteocephalus aggregate). In addition, G. osculata de Chambrier et al., 2004, 2015; Scholz et al, 2007). is much bigger and more robust than the species of the Species of both monotypic genera differ from each other Proteocephalus aggregate and its proglottids contain in their scolex and strobilar morphology, especially in the much more proglottids, especially immature ones, and presence/absence of a metascolex (absent in the former more testes (Scholz & Hanzelovâ, 1998; present study). species versus well-developed in P. parasiluri, which This tapeworm has been reported from the following was even placed in the subfamily Corallobothriinae - countries of Europe and Palaearctic Asia: Bulgaria, Freze, 1965), the position of bands of vitelline follicles Czech Republic, Germany, Hungary, Latvia, Lithuania, (‘typically’ vertical alongside margins of proglottids Poland, Russia, Slovakia, Sweden, Ukraine, United Kingdom, former Yugoslavia, Azerbaidzhan, Georgia, Iraq and Turkey (Fig. 16; see also Scholz & Hanzelovâ, 1998 and Gibson et al, 2005 [but misidentification of Postgangesia marmata de Chambrier, Al-Kallak & Mariaux, 2003 and Postgangesia hemispherous (Rahemo & Al-Niaeemi, 2001) in Iraq cannot be excluded], Glanitaenia osculata is a specific parasite of weis catfish, which is currently considered to be an invasive species and has recently been introduced to the western and southwestern Europe including northern Italy, Spain, France, Germany and England (Copp et al, 2009; Fig. 16). Therefore, it is possible that new geographical records from two of these countries (the River Po in Italy and the River Rhine basin in Germany - River Aare is a tributary of the Rhine joining it in Koblenz, at the Swiss- German border) are a result of this recent expansion of the weis catfish throughout Europe (Fig. 16). ACKNOWLEDGEMENTS Alexander Brinker, Fisheries Research Station of Baden Württemberg, Langenargen, Germany, Bahram S. Dezfuli, Fig 16. Geographical distribution of Glanitaenia osculata University of Ferrara, Italy, and Petr Horâk, Charles (Goeze, 1782) in Europe. Countries with previous University in Prague, Czech Republic, kindly provided records indicated by triangles; new geographical specimens of Glanitaenia osculata from Germany, records (Italy, Romania and Switzerland) highlighted Italy and Romania, respectively. The authors thank by asterisk); the original distribution area of weis Janik Pralong, Gilles Roth and André Piuz for technical catfish (grey silhouette; bordered by dashed line) and assistance, help with drawing and SEM micrographs, newly colonized regions of Europe (white silhouette; bordered by dotted line; modified from Copp et al, respectively, Jacqueline Studer and Julien Oppliger 2009). (all Natural History Museum Geneva, Switzerland) for Amended diagnosis of Glanitaenia 9 providing fish for parasitological examination, Roman phalata in Fish, Amphibians and Reptiles. Izdatel’stvo Kuchta and Blanka Skorikovâ Institute of Parasitology, “Nauka ”, Moskva, 538 pp. (In Russian: English translation, Ceské Budëjovice, Czech Republic, for preparation of Israel Program of Scientific Translation, 1969, Cat. No. 1853. v +597 pp). Fig. 16 and help with preparation of plate with Figs 1-9, Gibson D.I., Bray R.A., Harris E.A. (Compilers) 2005. 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