identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
744587ACFFC2D52E9D2C53C9FC540780.text	744587ACFFC2D52E9D2C53C9FC540780.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Unio tumidus Retzius 1788	<div><p>The U. tumidus lineage</p> <p>The U.tumidus lineage included samples from European rivers (namely, the Seine, Danube, Rhine, Thames and Weser), a Swedish lake, the Izorka and Kovash rivers in Russia and the Dnieper River in Ukraine (Fig. 2). This species is absent from the Iberian Peninsula and North Africa. Samples from the Rhine and Thames rivers and Bjornsjon Lake (Sweden) shared the same haplotype (Supporting Information, Fig. S3).</p> <p>The species had an elongated shell with an anterior umbo, but a characteristic cuneiform posterior end with a descending dorsal posterior margin, which clearly distinguishes this taxon from the other species. Nevertheless, there were specimens with short shells that were very difficult to identify (Fig. 3). We did not have access to the Ukrainian and Russian shells.</p></div> 	http://treatment.plazi.org/id/744587ACFFC2D52E9D2C53C9FC540780	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Araujo, Rafael;Buckley, David;Nagel, Karl-Otto;García-Jiménez, Ricardo;Machordom, Annie	Araujo, Rafael, Buckley, David, Nagel, Karl-Otto, García-Jiménez, Ricardo, Machordom, Annie (2018): Species boundaries, geographic distribution and evolutionary history of the Western Palaearctic freshwater mussels Unio (Bivalvia: Unionidae). Zoological Journal of the Linnean Society 182 (2): 275-299, DOI: 10.1093/zoolinnean/zlx039, URL: https://academic.oup.com/zoolinnean/article/182/2/275/4091584
744587ACFFC2D52E9EB451D1FABC0567.text	744587ACFFC2D52E9EB451D1FABC0567.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Unio durieui Deshayes 1847	<div><p>The U. durieui + U. gibbus lineage</p> <p>This clade grouped the samples from the morphospecies U. durieui and U. gibbus as sister lineages (Fig. 1). The U. durieui lineage only included samples from Tunisia and consisted of five different haplotypes among the specimens from the three rivers sampled. Distinct haplotypes were present in each river, but the most frequent haplotype was shared among specimens from the Ziatine and El Maaden rivers (Supporting Information, Fig. S1).</p> <p>Morphologically, U. durieui specimens were elongated with an intermediate shape between U. mancus and U. pictorum, and probably impossible to distinguish in the absence of geographical information. There were also rounded morphotypes similar to U. gibbus, as in the case of El Maaden (Khalloufi et al., 2011: fig. 5). This species was not found in Morocco but is present in Tunisia and Algeria to the east of the Moulouya River (Fig. 2).</p> <p>For the morphospecies U. gibbus, we obtained two clearly separated groups (genetic distance of 2.03%, Table 1): one representing haplotypes from the Iberian Peninsula (Barbate River) and Morocco, and another consisting of Tunisian haplotypes (Fig. 2; Supporting Information, Fig. S2). These two subclades were consistently recovered as distinct species in the bGMYC and M-PTP analyses. From 16 to 22 substitutions separated the Spanish-Moroccan haplotypes from the Tunisian ones. Although the eastern haplotypes were more similar, none of the Spanish, Moroccan and Tunisian specimens shared haplotypes among the 10 detected.</p> <p>This species had a rounded shape similar to Potomida, but clear diagnostic characters such as the lateral teeth, the umbonal sculpture and shell thickness differentiate the two taxa (Araujo et al., 2009a: fig. 2).</p> </div>	http://treatment.plazi.org/id/744587ACFFC2D52E9EB451D1FABC0567	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Araujo, Rafael;Buckley, David;Nagel, Karl-Otto;García-Jiménez, Ricardo;Machordom, Annie	Araujo, Rafael, Buckley, David, Nagel, Karl-Otto, García-Jiménez, Ricardo, Machordom, Annie (2018): Species boundaries, geographic distribution and evolutionary history of the Western Palaearctic freshwater mussels Unio (Bivalvia: Unionidae). Zoological Journal of the Linnean Society 182 (2): 275-299, DOI: 10.1093/zoolinnean/zlx039, URL: https://academic.oup.com/zoolinnean/article/182/2/275/4091584
744587ACFFC2D5209D2C51ADFBFE0735.text	744587ACFFC2D5209D2C51ADFBFE0735.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Unio foucauldianus Pallary 1936	<div><p>The (U. foucauldianus + U. delphinus) + U. tigridis, U. ravoisieri, U. mancus, U. elongatulus, U. pictorum lineage</p> <p>This lineage was the most complex within the genus (Fig. 1). All morphospecies were recovered as monophyletic groups, although some had high levels of within-clade diversity and phylogenetic structuring. The relationships among lineages, however, were not well supported.</p> <p>The first well-supported clade of this lineage included two closely related morphospecies with a genetic distance of 3.32% (Fig. 1; Table 1): U. delphinus from Atlantic Iberia and U. foucauldianus from Atlantic and Mediterranean Morocco (Fig. 2). The network analysis showed a minimum of 24 steps separating the two species (Supporting Information, Fig. S4). There was greater Atlantic/South differentiation in U. delphinus (no shared haplotypes between the 2 regions and up to 24 substitutions among the most differentiated haplotypes) than in U. foucauldianus (in this case, the main haplotype was found in both the Atlantic and Mediterranean basins).</p> <p>Many of the U. delphinus and all of the U. foucauldianus specimens from Mediterranean Morocco had an elongated shell with a pictorum shape, while specimens from the Atlantic rivers of Morocco and southern Spain were more rounded, resembling the mancus shape (Figs 4, 5).</p> <p>The second clade comprised U. tigridis samples from Israel (Lake Kinneret in the Jordan River basin) and Turkey (the Tersakan stream) (Fig. 2). The M-PTP analysis considered these samples as two different species (Fig. 1), with a genetic distance of 2.89% (Table 1). Only six specimens were analysed from this area, and the two haplotypes detected were separated by 32 substitutions (Supporting Information, Fig. S5).</p> <p>This species is typified by a short shell with anteriorly shifted umbos, which was observed in the Israeli specimens. The Turkish specimens resembled the U. mancus morph (Fig. 6).</p> <p>The next lineage combined the U. ravoisieri specimens into two subclades with a genetic distance of 2.55% (Table 1; Fig. 2). These subclades were also consistently recovered as distinct species in the bGMYC and M-PTP analyses. The two most frequent haplotypes (Supporting Information, Fig. S6) were separated by 18 steps. One of these haplotypes represented the Spanish samples, and was very close to a haplotype present in the Tunisian Kebir River (separated by only one step), and to one of the two divergent El Maaden haplotypes (separated by four steps).</p> <p>The shell of this species typically has a modified U. mancus shape, but the morphs we obtained were highly variable, especially those from the Spanish Ser River (Khalloufi et al., 2011: fig. 3).</p> <p>The clade consisting of U. mancus (Fig. 1) included specimens from Mediterranean basins in Spain (the</p> <p>The intraspecific divergences within species divided into different units were Unio crassus: 2.10, Unio pictorum: 0.72, Unio ravoisieri: 1.36, Unio tigridis: 0.96 and Unio gibbus: 1.18%.</p> <p>Ebro, Fluviá, Ter, Sonella, Llobregat and Júcar rivers), France (the Hérault and Argens rivers, the Tet River in the Basse Basin, Lake Bourget and the Ognon River, both within the Rhône Basin, many rivers in Corsica and the Atlantic rivers Charente, Seine and Drée, a tributary of the Loire) and Italy (many rivers in Sardinia) (Fig. 2). The network analysis showed haplotypes with some biogeographic incongruences (Supporting Information, Fig. S7). For instance, the Júcar and Sonella rivers (eastern Spain) shared a haplotype with some French populations, but not with any of the northeastern Spanish populations. This high-frequency haplotype appeared in both Atlantic and Mediterranean French and Spanish rivers. Of the 15 different haplotypes found, five were exclusive to Sardinian-Corsican populations, one to the French Lake Bourget and one appeared with a high frequency in the Catalonian (eastern Spain) rivers.</p> <p>This species had a more rounded shell than the other Mediterranean species U. elongatulus, with the exception of specimens from northeastern Spain, Corsica and Sardinia, which were elongated (Figs 7, 8).</p> <p>The next clade included the U. elongatulus (Fig. 1) specimens from Mediterranean freshwaters in Italy (Po and Isonzo rivers and Alpine lakes), Croatia (Mirna and Zrmanja rivers, Bačinska Lake) and Albania (Scutari Lake) (Fig. 2). The Croatian haplotypes appeared at both ends of the network (Supporting Information, Fig. S8), therefore, shared haplotypes between Croatia and Italy would be expected. However, no haplotypes were shared among any of the specimens from these regions.</p> <p>The specimens had more elongated shells than other Mediterranean species (U. mancus) and a posterior umbo, although some populations had more rounded shells (Fig. 9).</p> <p>The last clade represented the species U. pictorum, which included two groups having a genetic distance of 2.38% (Table 1). The first group (1) consisted of specimens from Western and Central Europe (Lake Bourget in the Rhône Basin and the Adour, Seine, Oder, Danube and Thames basins), eastern Greece (Strymonas and Axios rivers and Lake Volvi), Ukraine (Teteriv at the Black Sea), Iran and Russia (Dzhankhot River at the Black Sea, Vyborg River in Karelia). The second group (2) comprised specimens from both western (Trichonida Lake at the Acheloos basin) and eastern (Axios River) Greece (Fig. 2). These two subclades were consistently recovered as distinct species in the bGMYC and M-PTP analyses. Up to 16 different haplotypes (Supporting Information, Fig. S9), with from one to a maximum of 23 substitutions, were found among the specimens of this species. There was a maximum of 8 steps of differentiation within group ‘1’ and 17 between two of the three ‘2’ haplotypes. The network shape of group ‘1’ represented polymorphic populations with two relatively frequent haplotypes, one of which was only found in specimens from the Thames River and the other shared by samples as far away as Iran and Greece. Haplotypes found in the Danube and Dzhankhot rivers and in Lake Volvi seemed to derive from the latter one.</p> <p>This highly polymorphic species had a typical elongated shell. However, some specimens were very different, for example those from Strymonas River, Lake Volvi and the Axios River (within both clades). Some populations resembled U. mancus (Lake Volvi, Thichonida and Strymonas rivers) (Figs 10, 11).</p> </div>	http://treatment.plazi.org/id/744587ACFFC2D5209D2C51ADFBFE0735	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Araujo, Rafael;Buckley, David;Nagel, Karl-Otto;García-Jiménez, Ricardo;Machordom, Annie	Araujo, Rafael, Buckley, David, Nagel, Karl-Otto, García-Jiménez, Ricardo, Machordom, Annie (2018): Species boundaries, geographic distribution and evolutionary history of the Western Palaearctic freshwater mussels Unio (Bivalvia: Unionidae). Zoological Journal of the Linnean Society 182 (2): 275-299, DOI: 10.1093/zoolinnean/zlx039, URL: https://academic.oup.com/zoolinnean/article/182/2/275/4091584
744587ACFFCCD5259EB4513AFBEF04D1.text	744587ACFFCCD5259EB4513AFBEF04D1.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Unio tumidiformis Castro 1885	<div><p>The U. tumidiformis + [U. bruguierianus + (U. ionicus + U. crassus)] lineage</p> <p>This lineage included two monophyletic groups with a COI genetic distance greater than 7.85%: a western group that included the endemic Iberian species U. tumidiformis (Fig. 2) and a second group (supported by a posterior probability of 0.99 and bootstrap values of 79 for ML and 94 for MP) comprising three morphospecies with a complex geographic structure (Fig. 1; Table 1).</p> <p>Up to nine haplotypes were found among the samples analysed for U. tumidiformis (Supporting Information, Fig. S10). Two were exclusive to the Portuguese Sado River but were close to some Spanish haplotypes. The most frequent haplotype was found in the Mira and Guadiana rivers.</p> <p>Within the second group, U. bruguierianus from eastern Greece (the Axios and Pinios rivers) (Fig. 2) was the sister species of U. ionicus + U. crassus (genetic distance of 4.28–4.45%), although this relationship was not highly supported. Unio ionicus lives in Albania and the Acheloos River (including Lake Lysimacheia) in western Greece (Fig. 2) and was the sister group of U. crassus (genetic distance of 4.30%). Unio crassus presented five subclades with a minimum divergence of 2.16% (Table 1) and haplotypes separated by at least 19 steps (Fig. 12). These five subclades included (1) U. crassus courtillierii Hattemann, 1859 from France and one sample from Sweden (both sequences obtained from GenBank); (2) samples from the Sofaditikos (Pinios catchment), Aliakmon/Aliakmonas and Sperchios rivers in eastern Greece; (3) Central European samples from the Rhine, Danube, and Rhône catchments; (4) samples from the Lissos River (eastern Greece); and (5) samples from western Turkey (Fig. 2). This group had the most complexity, which is also reflected in the bGMYC and M-PTP analyses in which up to six species were revealed.</p> <p>All U. crassus haplotypes were represented in a unique network (Fig. 12). No clear biogeographic structure was observed, and some haplotypes belonging to specimens from the same rivers had a high number of substitutions (up to 33). From the right end of Figure 12, three haplotypes from eastern Greece were assigned to U. bruguierianus. Three distinct haplotypes from western Greece and Albania were considered to represent U. ionicus. Three haplotypes were found from the Turkish locations analysed, but with a smaller genetic distance compared with the other haplotypes.</p> <p>A polymorphic shell shape was observed for U. bruguierianus: some specimens from the Axios and Pinios rivers had a shell shape resembling some pictorum specimens from the Axios River, while other specimens from the Pinios River were identical to some U. crassus specimens from the Lissos River or Turkey (Fig. 13). This polymorphic shell shape was also observed for U. ionicus and U. crassus (Figs 13, 14). Unio ionicus had an almost identical shell to some U. crassus specimens from the Sofaditikos (Pinios Basin), Sperchios and Lissos rivers in Greece. In contrast, other U. crassus specimens from the Lissos River and from Turkey presented an elongated shell shape. There were also specimens with and without sculpture in the umbo area from the same locality with the same haplotype, such as those from the Lissos River (Fig. 14).</p> <p>The coalescence-based molecular clock analyses for the species tree resulted in very broad temporal intervals for most of the clades. Although the results have to be interpreted with caution, they provide a broad temporal framework for the diversification of the group. The coalescence reconstruction (Fig. 15) placed the common ancestor of the Unio clade and the origin of the Western Palaearctic Unio in the Eocene. In fact, the Western Palaearctic species underwent two main cladogenetic events. The first event occurred in the Early Eocene (U. gibbus, U. durieui split), and the second during the Oligocene, involving the divergence of the most speciose clade. Most of the modern species appeared during the Miocene. The most recent cladogenetic event involved the U. delphinus and U. foucauldianus clade at the end of the Miocene (Messinian). Notably, the reconstructed species tree agrees with the general topology of the concatenated matrix analyses except in the phylogenetic position of U. tumidus. In the species tree, U. tumidus appeared with strong support as sister group of the U. tigridis + [U. pictorum, U. ravoisieri (U. mancus + U. elongatulus, U. delphinus + U. foucauldianus)] lineage (Fig. 15).</p> </div>	http://treatment.plazi.org/id/744587ACFFCCD5259EB4513AFBEF04D1	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Araujo, Rafael;Buckley, David;Nagel, Karl-Otto;García-Jiménez, Ricardo;Machordom, Annie	Araujo, Rafael, Buckley, David, Nagel, Karl-Otto, García-Jiménez, Ricardo, Machordom, Annie (2018): Species boundaries, geographic distribution and evolutionary history of the Western Palaearctic freshwater mussels Unio (Bivalvia: Unionidae). Zoological Journal of the Linnean Society 182 (2): 275-299, DOI: 10.1093/zoolinnean/zlx039, URL: https://academic.oup.com/zoolinnean/article/182/2/275/4091584
