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55-518: Pelophylax is a genus of true frogs widespread in Eurasia, with a few species ranging into northern Africa. This genus was erected by Leopold Fitzinger in 1843 to accommodate the green frogs of the Old World , which he considered distinct from the brown pond frogs of Carl Linnaeus ' genus Rana . They are also known as water frogs , as they spend much of the summer living in aquatic habitat ;

110-556: A clade – that also includes such genera as Babina , Glandirana , Hylarana , Pulchrana , Sanguirana , Sylvirana , as well as Hydrophylax which like Pelophylax is suspected of being not monophyletic. These genera were formerly also included in Rana by most authors, and several of them have only been established in the 1990s. And as regards the possible paraphyly of Pelophylax , it seems that some species assigned there are very close to Hylarana , and thus it might simply be

165-557: A species : see Botanical name and Specific name (zoology) . The rules for the scientific names of organisms are laid down in the nomenclature codes , which allow each species a single unique name that, for animals (including protists ), plants (also including algae and fungi ) and prokaryotes ( bacteria and archaea ), is Latin and binomial in form; this contrasts with common or vernacular names , which are non-standardized, can be non-unique, and typically also vary by country and language of usage. Except for viruses ,

220-415: A complete ( haploid ) genome of only one parent species. Usually one entire genome of the parental species is excluded prior to meiosis during gametogenesis , such that only one (remaining) parental genome is represented among gametes and genes from the other parent are not passed on by the hybridogen. This discarding occurs gradually during subsequent mitotic divisions, not in one step. Hybridogenesis

275-416: A female P. ridibundus (RR). The offspring consist of P. kl. esculentus males (75%) or P. ridibundus females (25%). This is called hybrid-amphispermy. All- hybrid populations (E system , EE–system ) consist exclusively of P. kl. esculentus – diploid RE and triploid LLR or RRL hybrids. There are even known tetraploid LLRR hybrids. All-hybrid populations inhabit the entire range of

330-528: A hybrid coexisting (living in sympatry ) with one of the parental species required for its reproduction. P. kl. esculentus for example in the most frequent L-E system must mate with P. lessonae to produce new hybrids, in the R-E system with P. ridibundus . Because these hybrids depend on other taxa as sexual hosts to reproduce ("parasitize" on them sexually), they are kleptons ("kl." in scientific names ). The Pelophylax esculentus complex consists of

385-643: A later homonym of a validly published name is a nomen illegitimum or nom. illeg. ; for a full list refer to the International Code of Nomenclature for algae, fungi, and plants and the work cited above by Hawksworth, 2010. In place of the "valid taxon" in zoology, the nearest equivalent in botany is " correct name " or "current name" which can, again, differ or change with alternative taxonomic treatments or new information that results in previously accepted genera being combined or split. Prokaryote and virus codes of nomenclature also exist which serve as

440-434: A lineage of Raninae not particularly close to Rana . But it also turned out that these Eurasian green frogs might not form a monophyletic lineage. The sheer number of species involved in the group of Pelophylax and its closest relatives means that it will probably be some time until the definite circumscription of this genus is resolved. The Pelophylax frogs belong to a group of moderately advanced Raninae – possibly

495-621: A long time and redescribed as new by a range of subsequent workers, or if a range of genera previously considered separate taxa have subsequently been consolidated into one. For example, the World Register of Marine Species presently lists 8 genus-level synonyms for the sperm whale genus Physeter Linnaeus, 1758, and 13 for the bivalve genus Pecten O.F. Müller, 1776. Within the same kingdom, one generic name can apply to one genus only. However, many names have been assigned (usually unintentionally) to two or more different genera. For example,

550-577: A matter of moving them to that genus. But hybridogenic speciation is running rampant in the Old World green frogs, and this obfuscates the data gained from DNA sequence analyses. Pelophylax is a rather old and well-represented genus, with articulated fossils from Europe known as far back as the Early Oligocene . It has been theorized that Pelophylax originated in Asia no later than 5 million years before

605-409: A reference for designating currently accepted genus names as opposed to others which may be either reduced to synonymy, or, in the case of prokaryotes, relegated to a status of "names without standing in prokaryotic nomenclature". An available (zoological) or validly published (botanical) name that has been historically applied to a genus but is not regarded as the accepted (current/valid) name for

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660-427: A taxon; however, the names published in suppressed works are made unavailable via the relevant Opinion dealing with the work in question. In botany, similar concepts exist but with different labels. The botanical equivalent of zoology's "available name" is a validly published name . An invalidly published name is a nomen invalidum or nom. inval. ; a rejected name is a nomen rejiciendum or nom. rej. ;

715-455: A total of c. 520,000 published names (including synonyms) as at end 2019, increasing at some 2,500 published generic names per year. "Official" registers of taxon names at all ranks, including genera, exist for a few groups only such as viruses and prokaryotes, while for others there are compendia with no "official" standing such as Index Fungorum for fungi, Index Nominum Algarum and AlgaeBase for algae, Index Nominum Genericorum and

770-621: Is P. ridibundus specific and type B is P. lessonae -like (differs only by 0.3% from type C ). Most of P. kl. esculentus have C or D phenotype of the P. lessonae , not P. ridibundus mtDNA. Distribution of these phenotypes don't reflect exactly typical matting patterns. Mitochondria along with the mtDNA are inherited exclusively from the female. Since the primary hybridisations producing P. kl. esculentus occur between P. ridibundus females (large) and P. lessonae males (small) and later are maintained through backcrosses P. kl. esculentus females with P. lessonae males (L–E system ),

825-1060: Is a hemiclonal mode of reproduction — half of a hybrid genome is transmitted intact clonally from generation to generation ( R genome in the L-E system) — not recombined with a parental species genome (L here), while the other half (L) is transmitted sexually — obtained (replaced) each generation by sexual reproduction with a parental species ( sexual host , P. lessonae in the L-E system). There are at least three hybridogenetic species ( hybrids ) of water frogs in Europe – edible frog Pelophylax kl. esculentus , Graf's hybrid frog Pelophylax kl. grafi and Italian edible frog Pelophylax kl. hispanicus . Their mating patterns are classified into several hybridogenetic systems: (capital abbreviations below scientific names are genotypes ) All these hybrids contain genome of marsh frog P. ridibundus (R) and genome of second parental species (L, P or B). Most of above hybridogenic systems consist of

880-596: Is discouraged by both the International Code of Zoological Nomenclature and the International Code of Nomenclature for algae, fungi, and plants , there are some five thousand such names in use in more than one kingdom. For instance, A list of generic homonyms (with their authorities), including both available (validly published) and selected unavailable names, has been compiled by the Interim Register of Marine and Nonmarine Genera (IRMNG). The type genus forms

935-464: Is essentially a reverse form of the L–E system. Hybrids P. kl. esculentus (genotype RL) exclude here the P. ridibundus (R) or P. lessonae (L) genome in a 3:1 ratio and make mainly clonal P. lessonae (L), less P. ridibundus gametes (R). One frog produce either L or R gametes or a mixture of both. Their lineages are maintained through backcrosses of a male P. kl. esculentus (RL) with

990-935: Is most widespread hybridogenetic system. It is found in Western Europe . Hybrids P. kl. esculentus (genotype RL) exclude here the P. lessonae genome (L) and make exclusively clonal P. ridibundus gametes (R). In other words, edible frogs produce gametes of marsh frogs! Their lineages are maintained usually through backcrosses of a female P. kl. esculentus (RL) with a male P. lessonae (LL). The offspring consist of only P. kl. esculentus . P. kl. esculentus hybrids (RL) can mate also with each other, but only 3% of resulting tadpoles (RR) survive to sexual maturity (97% do not). The genomes of interhybrid crosses are female, because of carrying X chromosomes of females from primary hybridisation. The P. ridibundus – P. kl. esculentus (R–E , RE , ridibundus–esculentus ) system inhabits Eastern Europe . It

1045-409: Is sometimes assigned to Pelophylax , but must be considered a nomen oblitum : The following fossil species are also known: Genus The composition of a genus is determined by taxonomists . The standards for genus classification are not strictly codified, so different authorities often produce different classifications for genera. There are some general practices used, however, including

1100-460: Is somewhat arbitrary. Although all species within a genus are supposed to be "similar", there are no objective criteria for grouping species into genera. There is much debate among zoologists about whether enormous, species-rich genera should be maintained, as it is extremely difficult to come up with identification keys or even character sets that distinguish all species. Hence, many taxonomists argue in favor of breaking down large genera. For instance,

1155-474: Is the type species , and the generic name is permanently associated with the type specimen of its type species. Should the specimen turn out to be assignable to another genus, the generic name linked to it becomes a junior synonym and the remaining taxa in the former genus need to be reassessed. In zoological usage, taxonomic names, including those of genera, are classified as "available" or "unavailable". Available names are those published in accordance with

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1210-829: The P. perezi genome (P) and make exclusively clonal P. ridibundus gametes (R). Their lineages are maintained in so called P–G system through backcrosses of P. kl. grafi (PR) with P. perezi (PP). The primary hybridisation which originated Italian edible frog Pelophylax kl. hispanicus (genotype RB) was: Hybrids Pelophylax kl. hispanicus (RB) discard the P. bergeri genome (B) and make exclusively clonal P. ridibundus gametes (R). Their lineages are maintained in so called B–H system through backcrosses of P. kl. hispanicus (PR) with P. bergeri (BB). Matting patterns of hybridogenetic water frogs don't fit precisely known definitions of hybridogenesis: The Pelophylax kl. esculentus complex frogs have either of four phenotypes of mtDNA : Type A

1265-621: The International Code of Zoological Nomenclature ; the earliest such name for any taxon (for example, a genus) should then be selected as the " valid " (i.e., current or accepted) name for the taxon in question. Consequently, there will be more available names than valid names at any point in time; which names are currently in use depending on the judgement of taxonomists in either combining taxa described under multiple names, or splitting taxa which may bring available names previously treated as synonyms back into use. "Unavailable" names in zoology comprise names that either were not published according to

1320-799: The International Plant Names Index for plants in general, and ferns through angiosperms, respectively, and Nomenclator Zoologicus and the Index to Organism Names for zoological names. Totals for both "all names" and estimates for "accepted names" as held in the Interim Register of Marine and Nonmarine Genera (IRMNG) are broken down further in the publication by Rees et al., 2020 cited above. The accepted names estimates are as follows, broken down by kingdom: The cited ranges of uncertainty arise because IRMNG lists "uncertain" names (not researched therein) in addition to known "accepted" names;

1375-789: The hybrid taxon – edible frog P. kl. esculentus (genotype RL) and parental species – marsh frog P. ridibundus (RR) and pool frog P. lessonae (LL). Hybrids are females and males, which is unusual, because hybrids of other hybridogenic species are only females. The primary hybridisation originating P. kl. esculentus (genotype RL) is: It occurs between P. lessonae (LL) males and P. ridibundus (RR) females , because smaller P. lessonae males prefer larger females. The lineages of hybrids are maintained later through other matings, described below. P. lessonae and P. ridibundus have distinct habitat requirements and usually don't live together. The P. lessonae – P. kl. esculentus (L–E , LE , lessonae–esculentus ) system

1430-560: The ice ages , the population of the common ancestor of both parental species of the edible frog was split into two. These populations diverged, but remained genetically close enough to be able to create fertile hybrids . However, when diploid edible frogs mate with each other, their offspring are often malformed, so there are no pure populations of edible frogs unless some triploid individuals are present (the E system described above). Introduction of alien species belonging to water frog complex ( Pelophylax esculentus complex), for example,

1485-474: The lessonae genome (L) and generates gametes of the P. ridibundus (R). In other words, edible frogs produce gametes of marsh frogs. The hybrid populations are propagated, however, not by the above primary hybridisations, but predominantly by backcrosses with one of the parental species they coexist (live in sympatry ) with (see below in the middle). Since the hybridogenetic hybrids require another taxon as sexual host to reproduce, usually one of

1540-404: The platypus belongs to the genus Ornithorhynchus although George Shaw named it Platypus in 1799 (these two names are thus synonyms ) . However, the name Platypus had already been given to a group of ambrosia beetles by Johann Friedrich Wilhelm Herbst in 1793. A name that means two different things is a homonym . Since beetles and platypuses are both members of the kingdom Animalia,

1595-620: The E system. L, R – P. lessonae and P. ridibundus haploid genomes ; LL , RR – do not survive to sexual maturity; * females only (eggs); ** L gametes are produced by L R , but author doesn't write whether they take part in reproduction or not. It is not clear, whether the primary hybridisation which originated Graf's hybrid frog Pelophylax kl. grafi (genotype PR) was: Unlike P. perezi and Pelophylax kl. grafi , P. ridibundus and P. kl. esculentus do not belong to native fauna of Iberian Peninsula . Hybrids P. kl. grafi (PR) discard

1650-469: The French botanist Joseph Pitton de Tournefort (1656–1708) is considered "the founder of the modern concept of genera". The scientific name (or the scientific epithet) of a genus is also called the generic name ; in modern style guides and science, it is always capitalised. It plays a fundamental role in binomial nomenclature , the system of naming organisms , where it is combined with the scientific name of

1705-442: The base for higher taxonomic ranks, such as the family name Canidae ("Canids") based on Canis . However, this does not typically ascend more than one or two levels: the order to which dogs and wolves belong is Carnivora ("Carnivores"). The numbers of either accepted, or all published genus names is not known precisely; Rees et al., 2020 estimate that approximately 310,000 accepted names (valid taxa) may exist, out of

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1760-448: The brown frogs, in line with the tendency to place any frog similar in habitus to the common frog ( R. temporaria ) in Rana . That genus, in the loose circumscription, eventually became a sort of " wastebin taxon ". Around 2000, with molecular phylogenetic studies becoming commonplace, it was discovered that Fitzinger's assessment was correct after all – not only is Pelophylax an independent genus, but it does in fact belong to

1815-489: The doubled set (LL from LLR or RR from RRL here) are used to produce haploid gametes (L or R respectively), whereas the remaining ones may be excluded (R from LLR or L from RRL) is known as meiotic hybridogenesis. In one Slovakian population however, triploid males (LLR) and diploid LR females generate clonal LL and clonal R gametes respectively, instead of recombined L and clonal LR. P. lessonae (LL) and P. ridibundus (RR) offspring do not survive to sexual maturity in

1870-673: The earliest Oligocene of Chartres-de-Bretagne , France, which appears to be from the Pelophylax kl. esculentus hybrid complex. The species P. aquensis (formerly Rana aquensis ) is known from the Late Oligocene of southern France, and fossil species become more common during the Miocene. Including named klepta ( hybridogenic species), Pelophylax sensu lato contains 25 species: Named klepta ( hybridogenic species) of Pelophylax are: In addition, one species has been described that

1925-447: The earliest known fossils, and then dispersed west. It may have colonized Europe in the wake of a cooling/drying trend and the resulting Eocene-Oligocene extinction event , as part of an overall replacement of Europe's previously tropical frog fauna of African origin (such as the pyxicephalid Thaumastosaurus ) by a more temperate fauna of Asian origin. The oldest Pelophylax specimen is an articulated but headless specimen known from

1980-579: The expected mtDNA phenotype of P. kl. esculentus would be the phenotype of P. ridibundus . This unexpected phenotype distribution might be explained in such a way that most of P. kl. esculentus lineages might go through at least one backcross between P. kl. esculentus male with P. lessonae female. And such phenotype pattern suggests, that primary hybridisations are rare. The introgression of P. lessonae mtDNA in P. ridibundus (type B ) might be caused by matting between P. ridibundus and P. kl. esculentus having P. lessonae mtDNA. During

2035-446: The form "author, year" in zoology, and "standard abbreviated author name" in botany. Thus in the examples above, the genus Canis would be cited in full as " Canis Linnaeus, 1758" (zoological usage), while Hibiscus , also first established by Linnaeus but in 1753, is simply " Hibiscus L." (botanical usage). Each genus should have a designated type , although in practice there is a backlog of older names without one. In zoology, this

2090-727: The generic name (or its abbreviated form) still forms the leading portion of the scientific name, for example, Canis lupus lupus for the Eurasian wolf subspecies, or as a botanical example, Hibiscus arnottianus ssp. immaculatus . Also, as visible in the above examples, the Latinised portions of the scientific names of genera and their included species (and infraspecies, where applicable) are, by convention, written in italics . The scientific names of virus species are descriptive, not binomial in form, and may or may not incorporate an indication of their containing genus; for example,

2145-447: The genome is transmitted to the next generation clonally (not excluded unrecombined intact genome), and only the other half sexually (recombined genome of the sexual host), the hybridogenesis is a hemiclonal mode of reproduction. For example, the edible frog Pelophylax kl. esculentus (mostly RL genome), which is a hybridogenetic hybrid of the marsh frog P. ridibundus (RR) and the pool frog P. lessonae (LL), usually excludes

2200-432: The idea that a newly defined genus should fulfill these three criteria to be descriptively useful: Moreover, genera should be composed of phylogenetic units of the same kind as other (analogous) genera. The term "genus" comes from Latin genus , a noun form cognate with gignere ('to bear; to give birth to'). The Swedish taxonomist Carl Linnaeus popularized its use in his 1753 Species Plantarum , but

2255-628: The largest component, with 23,236 ± 5,379 accepted genus names, of which 20,845 ± 4,494 are angiosperms (superclass Angiospermae). By comparison, the 2018 annual edition of the Catalogue of Life (estimated >90% complete, for extant species in the main) contains currently 175,363 "accepted" genus names for 1,744,204 living and 59,284 extinct species, also including genus names only (no species) for some groups. The number of species in genera varies considerably among taxonomic groups. For instance, among (non-avian) reptiles , which have about 1180 genera,

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2310-406: The lizard genus Anolis has been suggested to be broken down into 8 or so different genera which would bring its ~400 species to smaller, more manageable subsets. Hybridogenesis in water frogs The fertile hybrids of European water frogs (genus Pelophylax ) reproduce by hybridogenesis ( hemiclonally ). This means that during gametogenesis , they discard the genome of one of

2365-403: The most (>300) have only 1 species, ~360 have between 2 and 4 species, 260 have 5–10 species, ~200 have 11–50 species, and only 27 genera have more than 50 species. However, some insect genera such as the bee genera Lasioglossum and Andrena have over 1000 species each. The largest flowering plant genus, Astragalus , contains over 3,000 species. Which species are assigned to a genus

2420-428: The name could not be used for both. Johann Friedrich Blumenbach published the replacement name Ornithorhynchus in 1800. However, a genus in one kingdom is allowed to bear a scientific name that is in use as a generic name (or the name of a taxon in another rank) in a kingdom that is governed by a different nomenclature code. Names with the same form but applying to different taxa are called "homonyms". Although this

2475-452: The parental species and produce gametes of the other parental species (containing a genome not recombined with the genome of the first parental species). The first parental genome is restored by fertilization of these gametes with gametes from the first species ( sexual host ). In all-hybrid populations of the edible frog Pelophylax kl. esculentus , however, triploid hybrids provide this missing genome. Because half of

2530-473: The parental species, they are called kleptons (with "kl." in scientific names ). There are three known hybridogenetic hybrids of the European water frogs: Hybridogenesis implies that gametes of hybrids don't contain mixed parental genomes , as normally occurs by independent chromosome segregation and crossover in meiosis (see also second Mendel's law , recombination ). Instead, each gamete carries

2585-517: The parental species. Because triploids discard this genome, which is available in one copy and leave two copies of second genome, they don't perform endoreduplication. Moreover, this not eliminated genome is transmitted to haploid gametes sexually , not clonally (recombined between two L's or between two R's), in contrast to the genome of diploid hybrids. Such modified hybridogenesis (or gametogenetic system ) occurring in allotriploid hybrids, where during meiosis chromosomes (genomes) from

2640-524: The pond frogs can be found more often, by comparison, on dry land, as long as there is sufficient humidity . Yet there are species of Eurasian green frogs – the Central Asian P. terentievi , or the Sahara frog ( P. saharicus ) – which inhabit waterholes in the desert . Most authors throughout the 19th and 20th century disagreed with Fitzinger's assessment. The green frogs were included again with

2695-526: The provisions of the ICZN Code, e.g., incorrect original or subsequent spellings, names published only in a thesis, and generic names published after 1930 with no type species indicated. According to "Glossary" section of the zoological Code, suppressed names (per published "Opinions" of the International Commission of Zoological Nomenclature) remain available but cannot be used as the valid name for

2750-497: The specific name particular to the wolf. A botanical example would be Hibiscus arnottianus , a particular species of the genus Hibiscus native to Hawaii. The specific name is written in lower-case and may be followed by subspecies names in zoology or a variety of infraspecific names in botany . When the generic name is already known from context, it may be shortened to its initial letter, for example, C. lupus in place of Canis lupus . Where species are further subdivided,

2805-412: The standard format for a species name comprises the generic name, indicating the genus to which the species belongs, followed by the specific epithet, which (within that genus) is unique to the species. For example, the gray wolf 's scientific name is Canis lupus , with Canis ( Latin for 'dog') being the generic name shared by the wolf's close relatives and lupus (Latin for 'wolf') being

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2860-403: The taxon is termed a synonym ; some authors also include unavailable names in lists of synonyms as well as available names, such as misspellings, names previously published without fulfilling all of the requirements of the relevant nomenclatural code, and rejected or suppressed names. A particular genus name may have zero to many synonyms, the latter case generally if the genus has been known for

2915-566: The values quoted are the mean of "accepted" names alone (all "uncertain" names treated as unaccepted) and "accepted + uncertain" names (all "uncertain" names treated as accepted), with the associated range of uncertainty indicating these two extremes. Within Animalia, the largest phylum is Arthropoda , with 151,697 ± 33,160 accepted genus names, of which 114,387 ± 27,654 are insects (class Insecta). Within Plantae, Tracheophyta (vascular plants) make up

2970-429: The virus species " Salmonid herpesvirus 1 ", " Salmonid herpesvirus 2 " and " Salmonid herpesvirus 3 " are all within the genus Salmonivirus ; however, the genus to which the species with the formal names " Everglades virus " and " Ross River virus " are assigned is Alphavirus . As with scientific names at other ranks, in all groups other than viruses, names of genera may be cited with their authorities, typically in

3025-547: The water frog complex. RL diploids discard L genome and produce gametes of P. ridibundus (R), or discard R or L genome and produce gametes of P. lessonae (L) or P. ridibundus (R) respectively. In both cases, diploid hybrids generate also not reduced diploid gametes (RL) needed to recreate triploids. Triploids LLR and RRL are providers of P. lessonae (L) and P. ridibundus gametes (R) respectively in this system lacking both of parental species. So triploid hybrids allow P. kl. esculentus populations to remain without

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