Introduction to Zoological Nomenclature

(Part 1)

Jerry Hooker, Department of PalaeontologyThe Natural History Musuem, London

TAXONOMY TAXONOMY

STABILITY OF BIOLOGICAL NOMENCLATURE STABILITY OF BIOLOGICAL NOMENCLATURE PROVIDES VITAL UNDERPINNING FOR TAXONOMY.PROVIDES VITAL UNDERPINNING FOR TAXONOMY.

Taxonomists must ensure that a given taxon can have only Taxonomists must ensure that a given taxon can have only one name by which it is properly known, thereby avoiding:one name by which it is properly known, thereby avoiding:

•Multiple names for one taxon: SYNONYMSMultiple names for one taxon: SYNONYMS

•The same name for multiple taxa: HOMONYMSThe same name for multiple taxa: HOMONYMS

This is accomplished through PRIORITY, TYPIFICATION This is accomplished through PRIORITY, TYPIFICATION and PUBLICATION. and PUBLICATION. Universality is achieved by the use of Latin for the Universality is achieved by the use of Latin for the construction of names.construction of names.

CODESCODESOriginally two, each ensuring stability of names in the two Originally two, each ensuring stability of names in the two divisions of life then recognized: divisions of life then recognized: Kingdoms PLANTAE and ANIMALIA. Kingdoms PLANTAE and ANIMALIA.

Later, the Bacteriological Code split from the Botanical Code Later, the Bacteriological Code split from the Botanical Code and a code of Virus nomenclature was established.and a code of Virus nomenclature was established.

NAME STARTING POINTSNAME STARTING POINTS 1ST CODE1ST CODEZOOLOGY:ZOOLOGY: 1st Jan. 1758 (Linnaeus)1st Jan. 1758 (Linnaeus) 19611961BOTANY:BOTANY: 1st May 1753 (Linnaeus)1st May 1753 (Linnaeus) 19061906BACTERIOLOGY:BACTERIOLOGY: 1st Jan. 19801st Jan. 1980 19581958

Athough the first comprehensive international codes long Athough the first comprehensive international codes long postdated the starting points in binominal nomenclature, their postdated the starting points in binominal nomenclature, their origins go back to Strickland, Darwin, Owen & Westwood origins go back to Strickland, Darwin, Owen & Westwood (1843) (Zoology) and de Candolle’s (1843) (Zoology) and de Candolle’s LoisLois of 1867 (Botany). of 1867 (Botany).

MODERN CLASSIFICATIONSMODERN CLASSIFICATIONSToday, thanks to cladistics, there are many more taxa of kingdom Today, thanks to cladistics, there are many more taxa of kingdom rank, grouped into three domains.rank, grouped into three domains.•Zoological CodeZoological Code covers: Metazoa; protistan taxa when workers covers: Metazoa; protistan taxa when workers treat them as animals for purposes of nomenclature; and treat them as animals for purposes of nomenclature; and ichnotaxa.ichnotaxa.•Botanical CodeBotanical Code covers: Plants (Embryophyta), Algae, protists covers: Plants (Embryophyta), Algae, protists regarded as plants, Fungi and Cyanobacteria.regarded as plants, Fungi and Cyanobacteria.

HOMONYMY BETWEEN CODES: OKHOMONYMY BETWEEN CODES: OK

Genus Genus FicusFicus - the fig tree - the fig tree Genus Genus FicusFicus - the fig shell - the fig shell

UNACCEPTABLE HOMONYMSUNACCEPTABLE HOMONYMS

TroglodytesTroglodytes Viellot, 1806 Viellot, 1806

TroglodytesTroglodytes É. Geoffroy É. Geoffroy Saint-Hilaire, 1812Saint-Hilaire, 1812

The earlier genus name for the wren is valid. The chimp is The earlier genus name for the wren is valid. The chimp is changed to the next oldest available name: changed to the next oldest available name: PanPan Oken, 1816. Oken, 1816.

SYNONYMSSYNONYMS

Law of priority determines that Law of priority determines that GlisGlis Brisson, 1762 rather Brisson, 1762 rather than than MyoxusMyoxus Zimmermann, 1780 is the valid genus Zimmermann, 1780 is the valid genus name for the edible dormouse.name for the edible dormouse.

BINOMINAL NOMENCLATURE FOR BINOMINAL NOMENCLATURE FOR THE SPECIESTHE SPECIES

Extract from Linnaeus (1758)Extract from Linnaeus (1758)10th edition of 10th edition of Systema NaturaeSystema NaturaeVol. 1 Regnum Animale.Vol. 1 Regnum Animale.

Genus Genus VespertilioVespertilio, with species, with speciesV. vampyrusV. vampyrus, , V. spectrumV. spectrum and andV. perspicillatusV. perspicillatus (bats). (bats).

Pre-Linnaean non-binominal Pre-Linnaean non-binominal names resembled diagnoses, names resembled diagnoses, so separation of the two was aso separation of the two was amilestone in taxonomy.milestone in taxonomy.

HIERARCHY - ZOO CODEHIERARCHY - ZOO CODE•Type of the family is the type genusType of the family is the type genus

•Type of the genus is the type speciesType of the genus is the type species

•Type of the species is the type specimen(s)Type of the species is the type specimen(s)

Such a system allows economy of diagnoses. Such a system allows economy of diagnoses. Thus, e.g. species A can be diagnosed as genus A Thus, e.g. species A can be diagnosed as genus A with the following characters….with the following characters….In the case of a monotypic nominal family or genus, In the case of a monotypic nominal family or genus, the diagnosis of the next rank down is the same as the diagnosis of the next rank down is the same as the nominal taxon.the nominal taxon.

Ranks above the family group (highest: superfamily) Ranks above the family group (highest: superfamily) and below the species group (lowest: subspecies) are and below the species group (lowest: subspecies) are not regulated by the Zoological Code.not regulated by the Zoological Code.

SUBSPECIES & SUBGENERASUBSPECIES & SUBGENERA

Gemmula (Gemmula) acutisinuata Gemmula (Gemmula) acutisinuata stubbingtonensis stubbingtonensis (Cossmann, 1899)(Cossmann, 1899)

Troglodytes troglodytes Troglodytes troglodytes troglodytestroglodytes (Linnaeus, 1758) (Linnaeus, 1758)

Trinominal or quadrinominal names resulting from use of Trinominal or quadrinominal names resulting from use of subspecies and/or subgenera do not affect the principle of subspecies and/or subgenera do not affect the principle of binominal nomenclature. The nominate subspecies takes its binominal nomenclature. The nominate subspecies takes its authorship and date from the species. Same procedure with authorship and date from the species. Same procedure with super- and sub- categories of genus and family.super- and sub- categories of genus and family.

RISKS OF INSTABILITYRISKS OF INSTABILITY

CuniculusCuniculus Brisson, 1762 Brisson, 1762

CuniculusCuniculus Meyer, 1790 Meyer, 1790

OryctolagusOryctolagus Lilljeborg, 1874 Lilljeborg, 1874

If Brisson (1762) is unavailableIf Brisson (1762) is unavailablethrough being inconsistentlythrough being inconsistentlybinominal, the replacement namebinominal, the replacement nameAgoutiAgouti Lacépède, 1799 must be Lacépède, 1799 must be used. The result is that the used. The result is that the established name for the rabbit,established name for the rabbit,OryctolagusOryctolagus, must be changed to , must be changed to the older namethe older name Cuniculus. Cuniculus.

PacaPaca

RabbitRabbit

Submissions are made to the ICZN and a vote Submissions are made to the ICZN and a vote results in anresults in an “Opinion”“Opinion” published in the published in the Bulletin Bulletin of Zoological Nomenclature.of Zoological Nomenclature.

E.g.E.g. Opinion 1894 Opinion 1894 in 1998 ruled that Brisson in 1998 ruled that Brisson (1762) should be rejected for nomenclatural (1762) should be rejected for nomenclatural purposes because it is not consistently purposes because it is not consistently binominal, but that certain genera from that work binominal, but that certain genera from that work should be conserved. These included both should be conserved. These included both CuniculusCuniculus and and GlisGlis..

PROBLEMS NOT SOLVED PROBLEMS NOT SOLVED BY THE CODE ALONEBY THE CODE ALONE

SECONDARY HOMONYMY OF SPECIESSECONDARY HOMONYMY OF SPECIES

When When MaurimontiaMaurimontia was synonymized with was synonymized withAiluravusAiluravus, A’s species name had to be , A’s species name had to be changed and became changed and became Ailuravus stehlinschaubiAiluravus stehlinschaubi Wood, 1976. Wood, 1976.

B. B. Ailuravus picteti Ailuravus picteti Rütimeyer, 1891Rütimeyer, 1891A. A. Maurimontia pictetiMaurimontia picteti Stehlin & Schaub, 1951 Stehlin & Schaub, 1951

If the genus name for If the genus name for Maurimontia pictetiMaurimontia picteti had had originallyoriginally been been AiluravusAiluravus, this would represent primary homonymy., this would represent primary homonymy.

OBJECTIVEOBJECTIVESYNONYMSSYNONYMS

SUBJECTIVESUBJECTIVESYNONYMSSYNONYMS

Platychoerops richardsoniiPlatychoerops richardsonii Charlesworth, 1855Charlesworth, 1855Miolophus planicepsMiolophus planiceps Owen, 1865 Owen, 1865based on same type specimen.based on same type specimen.

Cebochoerus minorCebochoerus minor Gervais, 1876Gervais, 1876based on different type based on different type specimens.specimens.

Cebochoerus helveticus Cebochoerus helveticus (Pictet (Pictet & Humbert,1869)& Humbert,1869)

N.B. These are minimal criteria and much more is expected now: N.B. These are minimal criteria and much more is expected now: primary type designation, a stand-alone diagnosis, differential primary type designation, a stand-alone diagnosis, differential diagnosis, type horizon & locality, thorough description, high diagnosis, type horizon & locality, thorough description, high quality illustrations, phylogenetic analysis, synonymy list etc.quality illustrations, phylogenetic analysis, synonymy list etc.

IMPORTANT DEFINITIONSIMPORTANT DEFINITIONSTo be an To be an AVAILABLE NAMEAVAILABLE NAME

Before 1931, it must have:Before 1931, it must have:•Been published after 1757Been published after 1757•Used Latin alphabetUsed Latin alphabet•Been consistently binominal if for a speciesBeen consistently binominal if for a species•Been valid for a taxon when first proposedBeen valid for a taxon when first proposed

These criteria constitute an These criteria constitute an INDICATIONINDICATION

After 1930, in addition it must:After 1930, in addition it must:•Have a description or definition purporting to Have a description or definition purporting to differentiate the taxon ordifferentiate the taxon or•Have a bibliographic reference to such orHave a bibliographic reference to such or•Be a replacement name Be a replacement name

VALID NAMEVALID NAME

A A VALID NAMEVALID NAME is an available name that is: is an available name that is:The correct name in an author’s taxonomic judgement The correct name in an author’s taxonomic judgement (i.e. not a junior synonym or a junior homonym), so long (i.e. not a junior synonym or a junior homonym), so long as it is code-compliant.as it is code-compliant.

A name that does not fulfill the criteria necessary to be A name that does not fulfill the criteria necessary to be an available name is:an available name is:A A NOMEN NUDUMNOMEN NUDUM

NOMEN OBLITUMNOMEN OBLITUM

NOMEN DUBIUMNOMEN DUBIUMA name of unknown or doubtful A name of unknown or doubtful application. E.g. if the type is lost application. E.g. if the type is lost or indeterminate. Names in or indeterminate. Names in common use that turn out to be common use that turn out to be nomina dubia require a Neotype. nomina dubia require a Neotype. Others are Others are best ignoredbest ignored and if and if unused since 1899 a nomen unused since 1899 a nomen dubium can become a:dubium can become a:

if applied from 2000 onwards. A if applied from 2000 onwards. A younger homonym or synonym younger homonym or synonym takes precedence, but a nomen takes precedence, but a nomen oblitum remains an available oblitum remains an available name.name.

PUBLICATIONPUBLICATIONCriteria:Criteria:•Issued for the purpose of providing a public and Issued for the purpose of providing a public and permanent scientific recordpermanent scientific record•Obtainable, when first issued, free or by purchaseObtainable, when first issued, free or by purchase•Produced in an edition containing simultaneously Produced in an edition containing simultaneously obtainable copies by a method that assures numerous obtainable copies by a method that assures numerous identical and durable copiesidentical and durable copies•Before 1986: on paper using conventional printing, Before 1986: on paper using conventional printing, hectographing or mimeographinghectographing or mimeographing•After 1985 & before 2000: includes other than After 1985 & before 2000: includes other than conventional printing providing other criteria are met.conventional printing providing other criteria are met.•After 1999: includes other than printing on paper After 1999: includes other than printing on paper provided it contains a statement that copies have been provided it contains a statement that copies have been deposited in art least 5 major publicly accessible deposited in art least 5 major publicly accessible libraries identified by name in the work itselflibraries identified by name in the work itself

NOT A PUBLICATIONNOT A PUBLICATION•When the work contains a disclaimer that it is not for public and When the work contains a disclaimer that it is not for public and permanent scientific record or for the purposes of zoological permanent scientific record or for the purposes of zoological nomenclaturenomenclature•Parts of a work where some or all of the names or nomenclatural Parts of a work where some or all of the names or nomenclatural acts are disclaimed for nomenclatural purposesacts are disclaimed for nomenclatural purposes•Published works suppressed by the Commission using its plenary Published works suppressed by the Commission using its plenary power, except as a source of descriptions & illustrationspower, except as a source of descriptions & illustrations•After 1930 handwriting reproduced in facsimileAfter 1930 handwriting reproduced in facsimile•Photographs as suchPhotographs as such•Proof sheetsProof sheets•MicrofilmsMicrofilms•Acoustic records as suchAcoustic records as such•Labels on specimensLabels on specimens•Copies of an unpublished workCopies of an unpublished work•Text or illustrations distributed by means of electronic signals Text or illustrations distributed by means of electronic signals •Abstracts issued primarily to participants at meetingsAbstracts issued primarily to participants at meetings

‘ ‘ WORKS, NAMES & NOMENCLATURAL WORKS, NAMES & NOMENCLATURAL ACTS’ (1999 Zoo code, p.123)ACTS’ (1999 Zoo code, p.123)

TYPE SPECIMENS 1TYPE SPECIMENS 1PRIMARY TYPESPRIMARY TYPES•HolotypeHolotype - original single specimen either designated or - original single specimen either designated or because there were no other specimens (monotypy).because there were no other specimens (monotypy).•SyntypeSyntype - original multiple specimens either designated; or all - original multiple specimens either designated; or all the material interpreted to have been used by an author in the the material interpreted to have been used by an author in the definition of a species, but where no types were designated.definition of a species, but where no types were designated.•LectotypeLectotype - single specimen selected subsequently from a - single specimen selected subsequently from a syntype series to resolve confusion.syntype series to resolve confusion.•NeotypeNeotype - single specimen designated subsequently to - single specimen designated subsequently to replace a missing holotype or lectotype or previously replace a missing holotype or lectotype or previously designated neotype or all the syntypes, specifically to resolve a designated neotype or all the syntypes, specifically to resolve a confusion.confusion.•Hapantotype Hapantotype - original single or multiple preparations or - original single or multiple preparations or cultures of an extant protistan, which cannot be restricted by cultures of an extant protistan, which cannot be restricted by lectotype selection, although some may be excluded if more lectotype selection, although some may be excluded if more than one species is found to be included.than one species is found to be included.

TYPE SPECIMENS 2TYPE SPECIMENS 2““SECONDARY” TYPESSECONDARY” TYPES•Paratype Paratype - Each specimen of a type series other than the - Each specimen of a type series other than the holotype.holotype.•ParalectotypeParalectotype - Each specimen of a former syntype series - Each specimen of a former syntype series remaining after selection of a lectotype.remaining after selection of a lectotype.OTHER TYPES, NOT REGULATED BY THE CODEOTHER TYPES, NOT REGULATED BY THE CODE•TopotypeTopotype - specimen from the type locality of the species - specimen from the type locality of the species or subspecies with which it is identified. Important for or subspecies with which it is identified. Important for judging variation where primary type is a single specimen, judging variation where primary type is a single specimen, and as a potential neotype.and as a potential neotype.•AllotypeAllotype - A designated specimen of opposite sex to the - A designated specimen of opposite sex to the holotype.holotype.•PlastotypePlastotype - cast of a type that is lost. May obviate the - cast of a type that is lost. May obviate the need for a neotype.need for a neotype.ABANDONED TYPES, SOMETIMES WRITTEN ON LABELSABANDONED TYPES, SOMETIMES WRITTEN ON LABELS•CotypeCotype - formerly used for either syntype or paratype. - formerly used for either syntype or paratype.•GenotypeGenotype - replaced by the term “type species”. - replaced by the term “type species”.

TYPE ACTIONS TYPE ACTIONS HOLOTYPE +/- PARATYPE(S)HOLOTYPE +/- PARATYPE(S)= stability= stability

SYNTYPESSYNTYPES= potential instability= potential instability

LECTOTYPE +/- PARALECTOTYPESLECTOTYPE +/- PARALECTOTYPES= stability= stability

If loss of primary type(s) means confusion:If loss of primary type(s) means confusion:

NEOTYPENEOTYPE

RECOGNIZING TYPESRECOGNIZING TYPES

You can’t necessarily rely You can’t necessarily rely on a statement in a on a statement in a publication or on a label publication or on a label that a given specimen is that a given specimen is a type.a type.

E.g. All specimens in E.g. All specimens in Paris figured by Cuvier in Paris figured by Cuvier in Ossemens FossilesOssemens Fossiles are are labelled “TYPE”. Some labelled “TYPE”. Some are, but can only be are, but can only be recognized by reference recognized by reference to articles in the Code & to articles in the Code & the relevant publications. the relevant publications.

SCOPE OF SYNTYPES: WIDESCOPE OF SYNTYPES: WIDEDepéret (1906) named Depéret (1906) named Catodontherium robiacenseCatodontherium robiacense from the localities Robiac & from the localities Robiac & Mormont. The only specimens Mormont. The only specimens he mentioned were from he mentioned were from Mormont previously fig’d by Mormont previously fig’d by Pictet & Humbert under Pictet & Humbert under another name. Sudre (1969) another name. Sudre (1969) selected a LECTOTYPE from selected a LECTOTYPE from Robiac that had never been Robiac that had never been figured but was assumed to figured but was assumed to have been seen by Depéret have been seen by Depéret when he named the taxon.when he named the taxon.Depéret’s syntypes included all Depéret’s syntypes included all the specimens that he could the specimens that he could have used to found his have used to found his species.species.

Pictet & Humbert 1869Pictet & Humbert 1869

Sudre 1969Sudre 1969

LOST TYPESLOST TYPESLophiotherium siderolithicumLophiotherium siderolithicum (Pictet, 1857)(Pictet, 1857)Holotype by monotypyHolotype by monotypyMISSING.MISSING.

L. siderolithicumL. siderolithicum from same from same horizon & locality as holotype:horizon & locality as holotype:Topotype - potential neotype, Topotype - potential neotype, but neotype designation but neotype designation unnecessary as figure of the unnecessary as figure of the holotype is adequate and holotype is adequate and consistent with existing consistent with existing topotypes.topotypes.

HONEST COMPOSITES HONEST COMPOSITES

The Syntype feet above are all composites according to The Syntype feet above are all composites according to the author, so each bone is a separate specimen. Multiple the author, so each bone is a separate specimen. Multiple taxa are involved so one bone needs to be selected as taxa are involved so one bone needs to be selected as Lectotype to stabilize the species.Lectotype to stabilize the species.

Eurytherium latipes Gervais, 1852

CLANDESTINE COMPOSITES CLANDESTINE COMPOSITES Owen’s fig.8 of one of his Owen’s fig.8 of one of his apparent Syntypes of apparent Syntypes of Thylacinus major Thylacinus major appears to be appears to be a chimaera, with characters of a chimaera, with characters of the genera the genera ThylacinusThylacinus and and SarcophilusSarcophilus. Examination of . Examination of specimens studied by Owen specimens studied by Owen shows that the outline of shows that the outline of specimen “B” matches cracks in specimen “B” matches cracks in the drawing. Fig.8 is therefore a the drawing. Fig.8 is therefore a composite drawing of teeth of composite drawing of teeth of Sarcophilus Sarcophilus (like A), a fragment (like A), a fragment of of Thylacinus Thylacinus jaw (B), & jaw (B), & probably a modern probably a modern ThylacinusThylacinus jaw. B is the Syntype as the jaw. B is the Syntype as the “teeth” specimen cannot be “teeth” specimen cannot be recognised. recognised. AA BB

88

Owen (1877)Owen (1877)

HOLOTYPE OR SYNTYPES?HOLOTYPE OR SYNTYPES?

The dispersed skeleton on the slab is holotype by monotypy. The dispersed skeleton on the slab is holotype by monotypy. Lectotype selection is invalid if only 1 individual in total.Lectotype selection is invalid if only 1 individual in total.

““Lectotype”Lectotype”

TYPES & VARIATIONTYPES & VARIATION

Treposciurus mutabilisTreposciurus mutabilis Schmidt-Kittler, 1970Schmidt-Kittler, 1970

Treposciurus gardneriTreposciurus gardneri Hooker, 1991Hooker, 1991

Treposciurus Treposciurus preeceipreecei Hooker, 1986Hooker, 1986

Variant like Variant like T. gardneriT. gardneri

Variant like Variant like T. mutabilisT. mutabilis

T. preeceiT. preecei based on designated syntypes to reflect polymorphism based on designated syntypes to reflect polymorphism and overcome confusion with the other two species.and overcome confusion with the other two species.

JUSTIFIED SUBSEQUENT EMENDATIONSJUSTIFIED SUBSEQUENT EMENDATIONS

Microchoerus wardiMicrochoerus wardi named after 2 people. named after 2 people. Emended to Emended to M. wardorumM. wardorum..

Plesiarctomys hurzeleriPlesiarctomys hurzeleri named after Hürzeler. named after Hürzeler. Emended to Emended to P. huerzeleriP. huerzeleri as named before 1985.as named before 1985.

CONCORDANCE OF ADJECTIVAL CONCORDANCE OF ADJECTIVAL SPECIES NAMES WITH THE GENUSSPECIES NAMES WITH THE GENUS

Palaeotherium minPalaeotherium minusus Cuvier, 1804 [neuter genus] Cuvier, 1804 [neuter genus]

Recombined as:Recombined as:

Plagiolophus minPlagiolophus minor or (Cuvier, 1804) [masculine genus](Cuvier, 1804) [masculine genus]

HYPHEN IN SPECIES NAME BANNEDHYPHEN IN SPECIES NAME BANNEDexcept:except:

““If the first element is a Latin letter used to denote If the first element is a Latin letter used to denote descriptively a character of the taxon..”descriptively a character of the taxon..”E.g. white letter hairstreak - E.g. white letter hairstreak - Strymonidia w-albumStrymonidia w-album..

DOMESTICATED ANIMALSDOMESTICATED ANIMALS

Equus caballusEquus caballus Linnaeus, 1758 Linnaeus, 1758Domestic horseDomestic horse

Equus ferusEquus ferus Boddaert, 1785 Boddaert, 1785Wild horseWild horse

Wild and domesticated equivalents do not enter into Wild and domesticated equivalents do not enter into synonymy. This is to avoid a domestic form taking priority synonymy. This is to avoid a domestic form taking priority over a wild form as would be the case here.over a wild form as would be the case here.

ICHNOTAXAICHNOTAXA

OphiomorphaOphiomorpha

CallianassaCallianassa

E.g.E.g.The ichnogenus The ichnogenus Ophiomorpha Ophiomorpha is is regarded as the fossil burrow of regarded as the fossil burrow of the ghost shrimp the ghost shrimp CallianassaCallianassa, but , but the names are not synonymsthe names are not synonyms..

1. Ichnotaxa must always be fossil.1. Ichnotaxa must always be fossil.

2. Homonymy with animals is not 2. Homonymy with animals is not allowed.allowed.

3. An ichnotaxon does not 3. An ichnotaxon does not compete in priority with a name compete in priority with a name established for an animal.established for an animal.

4. When establishing an 4. When establishing an ichnotaxon, the new name is ichnotaxon, the new name is followed by e.g. igen. et isp. followed by e.g. igen. et isp. nov., rather than gen. et sp. nov.nov., rather than gen. et sp. nov.

OPEN NOMENCLATUREOPEN NOMENCLATURE(Based mainly on recommendations by Bengtson (1988))(Based mainly on recommendations by Bengtson (1988))

•aff.aff.: relates a new, undescribed taxon to a named taxon; : relates a new, undescribed taxon to a named taxon; e.g. aff. e.g. aff. Agenus aspeciesAgenus aspecies (for a new genus), (for a new genus), Agenus Agenus aff. aff. aspecies aspecies (for a new species) and aff. (for a new species) and aff. AgenusAgenus aff. aff. aspecies aspecies (for new genus and species).(for new genus and species).•cf.cf.: (positioned as in “aff.”) identification is provisional.: (positioned as in “aff.”) identification is provisional.•??: identification is uncertain. It is in Roman script and follows : identification is uncertain. It is in Roman script and follows the name.the name.•sp. sp. oror subsp. subsp. (the latter clearer than ssp.):(the latter clearer than ssp.): specific or specific or subspecific identification impossible or not attempted.subspecific identification impossible or not attempted.•‘…’‘…’: obsolete name. Often used to denote paraphyletic taxon : obsolete name. Often used to denote paraphyletic taxon or one that is likely to be replaced following taxonomic or one that is likely to be replaced following taxonomic revision.revision.

Comment:Comment: In a database, one could infer simply the absence of In a database, one could infer simply the absence of a species or subspecies name or nothing below a higher rank to a species or subspecies name or nothing below a higher rank to indicate the level of identification achieved.indicate the level of identification achieved.

SYNONYMY LISTS SYNONYMY LISTS (from Matthews (1973))(from Matthews (1973))

Muscardinus avellanariuszzzzzzzzzzzzzzzzzzzzzzzzzzzzzzzMuscardinus avellanariuszzzzzzzzzzzzzzzzzzzzzzzzzzzzzzz