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This article isprotected Allrights bycopyright. reserved. doi: 10.1111/2041-210X.12594 thisversionandthe between lead todifferences been through the copyediting, typesetting, and proofreading which may pagination process, forpublicati accepted articlehasbeen Authors: Running : Title: IsaacEditor: Nick Article :Review Date:19-Apr-2016 Accepted Revised Date:08-Apr-2016 :07-Sep-2015 Received Date Accepteddatabases. However, thereno consensus is aboutONqualifiers andtheir meaning. biological disciplines, anda high amount of dataleft in ON can be found in literature and and abbreviations Open known Nomenclaturequalifiers. as (ON) Thiswidely approach is applied across 1. The uncertainty orthe provisionalstatus of ataxonomic identification can be expressed byaset of terms Abstract: Corresponding author: Ed Road, Wharf 647, Contees 2 Present address: Marine Invasion Research Laboratory,Smithsonian Environmental Research Center (SERC), Italy Venice, 2737/F, I-30122 - Mari ISMAR Article ofItaly, Council Research - National 1 CNR address: Authors’ Open in the biodiversity era era biodiversity the in Nomenclature Open MarcoSigovini

Review Nomenclature of Open

[email protected] [email protected] 1 , Keppel, Erica gewater, Maryland USA 21037 USA21037 Maryland gewater, 1,2 , Davide Tagliapietra on and undergone full peer review buthasnot review undergone fullpeer on and Version of Record. PleaseVersion cite thisarticle as ne Institute, Ar Institute, Sciences ne 1

senale Tesa 104, Castello 104,Castello Tesa senale This article isprotected Allrights bycopyright. reserved. since it cannot exactly, be definitely readily or This determined. uncertaintymayvarious from derive causes, be finally entered in databasessubsequent for elaboration. Sometimes the identity of aspecimen is uncertain burden torecognize andname each individual of aparticular assemblage andtocompile SA matrices thatwill still largely is identification and description 2011). Erwin & Hamilton (Pearson, the increasing need taxonomic expertise, of face howthe to decline in trained taxonomists is still debated reconstructingseries andbiodiversity trends with therecovering andanalysis of historical .Despite buildin in commitment aworldwide is there time, same number of dataare produced, generally in the of Species/Abundance(SA) matrices. Atthe Programs the in Australia, monitoring rely on of biological communities.consequence, As a anincreasing programs all over theincluding world, the National Estuary Program in the USA andthe Marine Monitoring assessment of water systems is largely based ontheir biodiversity andcommunity structure. Other large established therequirements for theecological quality of European the waters. thisframework, In the quality (2008/56/EC) Directive Framework Strategy Marine European the recently, more and, (2000/60/EC) biodiversity andenvironmental quality assessment. ago Adecade theFramework European Water Directory An increasing numbermonitoring of plans requires taxonomic identification for purposes the of both Keyword: offeringstarting a point for amethodological harmonization. biodiversity programs,well as for as researchers dealing with biodiversity datainfrastructures andtools, 3. This review provides aguide for taxonomists and identificati during ofuncertainties sources the clarify is recommendation . Some incons given toavoid 2. The use of ONqualifiers has been reviewed in toprovide asummary andguide to current practice in Accepted ArticleIntroduction tools, Species/Abundance matrices,, Taxonomic Databases, Uncertainty, Zoology , Biomonitoring, Controlled Biomonitoring, Biodiversity Informatics,

on and to facilitate the a tofacilitatethe and on based on morphological trai morphological on based ecologists currently involved in biomonitoring and and biomonitoring in involved currently ecologists istencies or vagueness. A flowchart isvagueness.istencies proposed A to or g large or even global- inventories, and in inventories,g large oreven global-scale andin Vocabulary, Biodiversity Primary Semantic Data, pplication of ON qualifiers. qualifiers. ofON pplication ts. The taxonomist has the the has taxonomist ts. The This article isprotected Allrights bycopyright. reserved. the design, implementation andmaintenance of biodiversity datainfrastructures andtools. who were notinvolved in the production of the data. Aconsistent and shared ONvocabulary would help with taxonomists workingidentifications, on butalso biodiversity informaticians,ecologistsand other datausers The harmonization of ONpractice would benefit people involved ateach step of biodiversity datalifecycle: 2005). (Chapman experience attribute tothe qualifiers the depends the , branch of laboratory on traditions andtheir own retains : when taxonomists orecologists produce orconsult taxonomic lists, the meaning they 1999 Winston 1936; McMasters & Schenk (e.g. procedures indica general Some 1988). Bengtson 1973; Matthews subject is inconsistent andlimited toahandful papers of predating the of digital information (e.g. ON qualifiers,since not form they do partof the nameof ataxon(Art. 5.3).The literature discussing the Internat The guidelines. or standards by uniform a for need the Despite . The procedure was developed primarily in palaeontology, essentially because of fragmentary the of al. their since diverged and asbotany such Ho 2003). Lester & Moulton (Stribling, Thewidely approach is applied across taxonomic disciplines, including bothproduction andresearch taxonomy uncertainty. not yet fully accomplished (Matthews 1973).ON qualifiersexpress may different kinds anddegrees of also called "qualifiers".Taxa are said tobe "left in open nomenclature" tounderline thatthe identification is terms andtheir abbreviations (called "signs"), which can be combined totaxon . Terms andsigns are of ensemble an on isbased approach The 1988). Bengston 1973; Matthews 1948; (Richter (ON) Nomenclature" express theuncertaintythe provisional or status of identification the using procedure a called "Open may belong toanon-indigenous species oraspecies unknown toscience. Inall these cases, scientistsmay specimen the case which in keys, identification the in species the of lack the to related be can it Moreover, envisagedmorphotypes different taxonomic or perspectiveswhich translates into identification resources. of orincompleteness stateofpreservation bad including

Accepted zoologicaldisciplines. in use ofON the on focused is paper This 2008). Article wever, nomenclature and notations from different biological branches branches biological different from notations and nomenclature wever, approach is felt for a long time(e.g isfeltforalong approach ional Code of Zoological Nomenclature Zoological of Code ional origin in both formal and substantive aspects (Minelli (Minelli aspects formaland substantive both in origin the specimen, variability of diagnostic characters, not not characters, ofdiagnostic variability specimen, the tions are presented in textbooks on taxonomic taxonomic on textbooks in presented are tions ). Nevertheless, the application of ON terms still still terms ofON application the ). Nevertheless, . Richter 1948), ON is not regulated isnot ON 1948), . Richter

(ICZN1999) does notdeal with et This article isprotected Allrights bycopyright. reserved. Acceptedspecies Information). All the qualifiers anddefinitions are summarized in alphabetical order in Appendix S1 (Supporting information. additional conveying time same the at resolution, taxonomic of level achieved the for different taxonviews. They also include qualifiers describing the stages ormorphotypes; they may account are presented in asubsequent . They include Moreover, other main qualifiersfrequently used in taxonomic records andclosely associated toONpractice 2005). (Chapman used alsobe may parentheses) between unique , such reference as a totheauthorandyear. Areferencevoucher to specimens(typically Articleliterature, describingmany different as just Therefore, taxa. provisional designations be should followed bya them (Matthews 1973).The combination of ataxonname and aqualifier may be found multiple in the and species names. the If author(and date) follows the taxonname, noqualifier should be inserted between no should signs tonames, attached When (1988). Bengtson with qualifiers ofthe placement The consistent. more is syntax ON the of use Current identification. the during arising uncertainties of sources main the summarize th Tofacilitate applied. inconsistently or vaguely currently are which terms those of use the regulate to given are advices Some meaning. similar drawn from aquatic extant . Some qualifiers are also used in other biological disciplines with the same or and discussed, in order toprovide asummary andguide tocurrent practice in zoology. All the examples are termsThemain andabbreviations ON found literature andother in the dataresources have been collected Annotated list of Open Nomenclature qualifiers , abbreviation: keys for the description ofaparticular morphotype. Otherwise, it could be aspecies notyet it instance for provisional; ispotentially uncertainty attributed todifferent causes (e.g. Matthews been has impasse this literature the In species. known any to related been has it nor level, species the to down identified been not has specimen the when generic the after used is qualifier This sp. (e.g.: (e.g.: e application of ON qualifiers, aflowch qualifiers, ofON e application sp.).

respect to the name name taxon tothe respect qualifiers expressing unresolved taxonomic status or 1973). According to the most common usage, the usage, common most the to According 1973). t be spelled in italics, which is reserved for forgenus isreserved italics,which in spelled t be could be could due tothesuitable lack dichotomous of art is proposedart is 1),which also (Fig. follows Matthews (1973) and and (1973) Matthews follows familia genusspecies genus species species This article isprotected Allrights bycopyright. reserved. Accepted Article

in the plural form, abbreviation: Nematoda sp.). Nematoda level.The qualifier also may be abbreviated unambig the to refers uncertainty the when isused it but "sp." of meaning same the has qualifier This rank, i.e. "sp." (e.g.: sp.). Zoarcidae follows thename of thefamily. The qualifiermay alsoabbreviated be unambiguously the lowest to genus level. Ithassame themeaning "sp." of butatahigher .The qualifier generally the at nor species the at neither identified been not has specimen the when isused qualifier This preferred. preferred. 1, sp. 2, etc). Conversely, in case diagnostic characters missing, are the qualifier "indet." should be specimens are analyzed, it should be possible toseparate them into different provisional species (sp. identificationwas notachieved. This qualifiershould be matrices, avoidedSAsince,when in integer whose genus, same the of species one than more of presence the describe to is used qualifier This "sp. A", "sp. B", etc. of them, the qualifiershould be followedsequential bya identifier,for instance"sp."sp. 1",etc. 2", or If distinctspecies recognized are within thegenus, same it is but notpossible toassignname a toany below). (see respectively qualifiers the use to we recommend cases these in uncertainty, us be itmay 1988).Since (Bengtson possible been of diagnostic characters,whenidentification or the not been attempted has despiteit could have lack or deterioration the to due indeterminable is specimen a when used been also has qualifier The belong tonon-indigenous orpreviously undescribedspecies. described. Therefore, specimens qualified with"sp." require careful consideration since they could , abbreviation: , abbreviation:

, abbreviation:

ssp gen. sp. . or subsp fam. gen. sp gen. fam. or . (e.g.: g. sp. spp.

, or (e.g.: gen. Zoarcidae sp.). trutta species plurimae . (e.g.: Nematoda fam. gen. sp.). sp.). fam.gen. Nematoda . (e.g.: Linnaeus, 1758 ssp.). 1758ssp.). Linnaeus, eful to distinguish among these sources of of sources these among todistinguish eful , abbreviation: uously tothelowest rank,i.e. "sp." (e.g.: species indeterminabilis species sp. pl. pl. sp. (e.g.: (e.g.: spp.). and stetit

abbreviation: species This article isprotected Allrights bycopyright. reserved. species nova sp.

Accepted (e.g.: Article

indeterminabilis the publication ( publication the Recom and Art.16.1 1999: (ICZN new intentionally of these qualifiers some strictly (or equivalent expression) toexplicitly indicate names taxa as circumstances. Onthe the one International , CodeZoological of Nomenclature requires the use different terms: slightly totwo totime from associated time The been "indet."forms andsimilar have abbreviation the same meaning "sp." of when the uncertainty refers tothe subspecific level. Subspecies is theonly rankbelowspecies the level ac sp.” (orasimilarexpression)constitutes act(e.g. anomenclatural qualifier can be also used athigher taxonomic ranks (e.g. indet.). mate caseofdamaged the in especially characters, indeterminable beyond acertain taxonomic level the due deterioration to of lack diagnostic or abbreviation of termsynonym ON of the as alsoused have been forms these both 2010).Occasionally, (e.g.Hawksworth, interchangeably meanings two the used have authors Some specimen. the of incompleteness or condition poor the indeterminabilis genus ( the subspecies ( in which the adjective is placed after the ,is The specimen is considered tobelong toanew, previously undescribed, species. Theform "sp. nov.", abbreviated species: with combination Tisbe or indet. sp. nov.). nov.). sp. nova species genus novum , ind. fam. nov. fam. , i.e. "species thatcannot be determined". The latter meaning is usually associated to , = indeterminable species (or species = indeterminable subspecies nova subspecies sp. indet. indeterminabilis species indeterminata = newspecies, abbreviation: , abbreviated ICZN 1999) of a binomen followed by an an by followed 1999) ICZN ofabinomen , fam. n. fam. species or gen. et sp. nov. sp. ind. , (e.g. Matthews 1973). We propose the "indet." to be used as an an as used be to "indet." sign the propose We 1973). Matthews (e.g. , abbreviated (as in Matthews 1973) andtoindicate (as inMatthews 1973) thatthe specimen is nov. fam gen. nov. gen. (e.g.: , i.e. "indeterminate, not determined species", and and species", determined not i.e."indeterminate, , . or , , nov. gen etsp. ssp. nov. ssp. g. nov. n. fam. species sp. nov. the most common. Similar terms are also applied to to applied also are terms Similar common. most the rial and missing parts (see e.g. Granzow 2000). This This 2000). Granzow e.g. (see parts rial and missing mendation 16A). When describing a new species, species, new a describing When 16A). mendation cepted by the ICZN (1999). As ON qualifier, it has ithas qualifier, ON As (1999). ICZN the by cepted , ). These qualifiers are used in two different different two in used are qualifiers These ). ,

gen. n. subsp. nov. indeterminata , indet.). spec. nov. spec. , etc.) and familia level ( level familia , etc.)and , abbreviation such as “sp. nov.” or“n. nov.” as“sp. such abbreviation g. n. benovici , , , ssp. n. ssp. sp. n. sp. nov. gen. = not determined= not species), , , nov. sp.

subsp. n.

, sp. nov. in Piraino nov. Piraino sp. in n. g. , nov. spec. , , etc.; also in familia nova n. ssp. n. , etc.), or n. , after the qualifier neither in full orabbr full in neither qualifier after the (e.g.: with" affinity "has for affinis This article isprotected Allrights bycopyright. reserved. Accepted Article , abbreviation: intended toindicate affinity, butin aweaker thanthe previous qualifiers, andis frequently 1936,isfalse McMasters & abbreviated species, implying thatit may belong toanew one. The term (abbreviated proxima species include These . in as such contexts, limited more in used been have qualifiers Similar affinis greater degree confidence of (Estes 1987). The qualifier is also found in literature the form in regard taxonomists However, other contrast in identity, possible a than more Be and (1986) to Lucas According 1988). (Bengston one, in terms of phylogenetic relatedness. Possibly, it may belong toanew, undescribed species of the species. Otherwise, it can be possibly attributed toa specieswith strong affinity with aknown desc species fromthe clearly differs specimen it. The Thisindicates qualifier thatthe specimen identical hassometo aknown to species affinity not butitis (e.g. a be avoided can by ambiguity any However, (1999). nov. This A). usage doesn’t imply anomenclatural is act and therefore inconsistent not with the ICZN follows the genus name is (and sometimes followed sequential bya identifier; e.g. In this case, the qualifier attached is toahigher-r 1988). Bengston 1973; (Matthews description formal publicationof the still taxabefore unnamed the On the other hand, they are commonly found in literature as ONqualifiers, as away to refer tonew, (2015). Cambefort et al. , 2014). The role of these qualifiers in nomenclatural texts is discussed in more detail by by detail more in discussed is texts nomenclatural in qualifiers ofthese role The , 2014). , abbreviated ex gr. ex aff. nr. or ; or ; or ("the nearest species", abbreviated abbreviated species", nearest ("the sp. A). or ex aff. sp. nr. sp. species affinis gr ., from the Latin

(e.g. Schenk & McMasters 1936), however this is a Latin misconstruction. misconstruction. isaLatin this however 1936), McMasters & Schenk (e.g. ), which indicate that the specimen is near but not identical to a known toaknown specimen butnotidentical thatthewhich indicate near is ), since the word the since etymology eviated form (Zidek 1987; Bengston 1988). 1988). 1987;Bengston (Zidek eviated form , abbreviation: them as synonymous (Zidek, 1987), or consider "aff." to indicate a to indicate "aff." consider 1987),or (Zidek, synonymous as them aff. alcocki to the meaning usually given tothe meaning usuallygiven term to the : herd, group; the form Koehler, 1914);it is notrequired torepeat the genus ank taxon: for instance, the abbreviation “sp. nov.” sp. aff. sp. ription, but it may fall within the variability limits limits variability the within fall itmay but ription, ngston (1998), the qualifier implies distinction distinction implies qualifier (1998),the ngston pplying a broader ON qualifier such as “sp. A” A” as“sp. such qualifier ON abroader pplying grupo prox. = affinity = with species, aknown from the does not exist in Latin) is also generally Latin) existin isalsogenerally not does or ex grege ex grupo sp. prox. sp. ("of the including", group , reportede.g. in Schenk ) and "species) near" confer Pristiophorus (see below). (see below). sp. sp. ex tubifex Latin confer species incerta 1988). genus the torepeat required This article isprotected Allrights bycopyright. reserved. Accepted Article confero , abbreviation: different approaches (Kornicker 1979). Following Following 1979). (Kornicker approaches different follow may name taxon the to respect with mark question the of placement the literature, the In in the identification with respect to Matth and (1948) Richter to According investigation. identification has been outcorrectly carried some but,forreason,still is uncertain needs and further unc of mark ("sign question The example in the ICZN (1999: Art. 11.9.3.4), we sugg 11.9.3.4),we Art. (1999: ICZN the in example meaning "similar", "of the same shape", as in Lucas (1986). (1986). Lucas asin shape", same "ofthe "similar", meaning for stand not does abbreviation The 1973). Matthews 1948, (Richter identification the with uncertainty" (see "sign of the synony them consider or 1987) (Estes view opposite to compared uncertainty with reference material consultingspecialist or a Itgenerally thetaxon. implies degree lower of a of comparing is tobe definitive likely after provisional is characters identification but The unclear. are This qualifier indicatesmost of that the diagnosticcharacters correspond toagivenspecies, butsome comma). is qualifier the cases, allthese In below). subjective. All they have beenfrequently used when dealing with species complexes groups and (see Pseudocandona import with groups microfaunal some in applied (infinitive (infinitive (Müller, 1774) sp. inc.). inc.). sp. 1774) (Müller,

= uncertain species, abbreviation: ' sp. nov. aff. aff. nov. sp. cf. , conferre exgr. cfr. , conf. after the qualifier neither in full or full in neither qualifier afterthe eremita ): "to bring together" (e.g.: (e.g.: "to bringtogether" ): or affinis rufescens sp. cf. sp. (Vejdovsky, 1882)). The differences between these terms are rather arerather terms these between differences The 1882)). (Vejdovsky, (Lucas 1987; Bengston 1998) however other taxonomists have the the have taxonomists other however 1998) Bengston 1987; (Lucas species incerta species = to compare, or tobe compared imperative =mood with, tocompare, from the confer ertainty", Matthews 1972) is attached to a taxon when the the when toataxon attached is 1972) Matthews ertainty", (Heller, 1861); the two qualifiers may also be separated by a by separated alsobe may qualifiers two the 1861); (Heller, , while Bengston (1988) consider them as almost . synonyms. asalmost them consider (1988) Bengston , while ? frequently used in combination with ', ', sp. inc. sp. Matthews (1973) and Bengston (1988), as well as an an as as well (1988), Bengston and (1973) Matthews ), but the latter should indicate a higher reliability in in reliability ahigher indicate latter should the ), but est toplace the sign the after taxonomic level to mous (Zidek, 1987). The term also overlaps in part part in alsooverlaps term The 1987). mous (Zidek, ews (1973), it should indicate a higher reliability reliability higher a indicate should it (1973), ews or (e.g.: such records as Ostracoda inc. cf. cf. abbreviated form (Zidek 1987; Bengston 1987;Bengston form(Zidek abbreviated (e.g.: (e.g.: hedgpethi Er. Marcus, 1964); it is not is not it Marcus, 1964); Er. (Müller, 1774) ? = = ? 1774) (Müller, species nova species conformis (e.g. , attempted, from the Latin for "he/she stood, stayed, remained" (e.g.: Teredinidae stet. = Teredinidae). Teredinidae). = stet. Teredinidae (e.g.: remained" stayed, stood, "he/she for Latin the from attempted, This article isprotected Allrights bycopyright. reserved. Accepted cetera animalia stetit Article , abbreviation: suggest using the term the using suggest furtheruseful, identification,even intothe go to To the best of ourknowledge, there is noqualifier in use toclearly state thatis notwanted, orfound also 1999;see (ICZN, uncertain is position taxonomic the that means which should notbe confused with digital form it may be conveniently substituted by"sp. inc.", "gen. inc.", etc. The term question mark. Since the latter may be considered as a"wildcard" bysomesoftware, in datastored in qualifiers tobe restricted tothe meaning of "uncertainidentification" andtobe equated tothe below). The abbreviations terms for the even 1973), and been usedwith different meanings, including as asynonym for "sp." and"sp. indet."(as Matthews in term The identifications. found in palaeontological studies, andis generally applied not during conventional taxonomic istypically approach This 1979). (Kornicker assignment entire the in or Art. 11.9.3.4), 1999: ICZN to species ofthe assignment the in uncertainty which it is applied. The question mark can be related to the genus level in abinomen toexpress It onscarce descriptive information, a Under this term are groupedall the unidentified specimens notlisted as separate taxain SA matrices. indet. may be merged into " with different ONqualifiers aunique into entry atasafe level (e.g. This approach can bealso usedwhensets,more aggregating datamerge records in is needed case to proceeding further.The intentional absence of anyqualifier should also be givenmeaning.same the mainly in technical reports. The term The reports. technical in mainly , abbreviation: species incerta species stet. (or intentional (or absence qualifier) any of =further identification has notbeen a.c. stetit (abbreviated as "sp. inc.") originally is unrelated tothe question mark. Ithas or gen. inc. gen. A.C. incertae after the taxon name to explicitly express the identifier choice of not not of choice identifier the express explicitly to name taxon the after species inquirenda = the remaining . animals. remaining = the and cetera " or" gen. et sp. inc. sp. et gen. ("of uncertain taxonomic position", abbreviated Corophium (abbreviated nd is usually employed withsummarizing purposes, established genera (Matthews 1973, Bengston 1988, 1988, 1973,Bengston (Matthews genera established and if allowed by the condition of the specimen. We We specimen. the of condition the by allowed if species dubia arealsofound. Wepropose usage the these of stet."). c. or

cet. ) may also be applied tohigh-rank , which are not ON qualifiers (see (see qualifiers ON arenot , which Corophium species inquirenda sp. and species incerta species Corophium inc. sed. inc. ). )

compass, infold" (e.g.: (e.g.: infold" compass, complex, species varietas forma Other qualifiers This article isprotected Allrights bycopyright. reserved. Accepted Article , abbreviation: , boundaries, typically due tophenotypical similarities. Itreferseither of toagroup sibling orcryptic The is generally understood as agroup of related species characterized by unclear or identical are reproductively that isolated" (Mayr 1963), although theterm also has by coined was species" "sibling term The species. in the sample, excluded). together c." listed "Peracarida (e.g. identified 45.6). 45.6). infrasubspeci identify "ab." and "morph" qualifiers the Aberrant specimens are pointed outby the qualifier Alternative growthforms orreproductive bemay recorded byadding specific qualifiers. mo as regarded more properly be should they (1999) ICZN the by regulated arenot subspecies of level the below categories since itin gave expressly author the unless subspecific, (with the exception of names published before 1961, which in case they havebe considered to In case of , thequalifier " used inconsistently andsometimes even interchangeably. common infrasubspecific categories among those introduced in the past; however have they been (McNeill and fungi, infrasubspecific categoriesexplicitly is regulated bythe International Code ofNomenclature for , Forma of meaning same the with taxa abbreviation: and

abbreviation: f. varietas = form (e.g.: =form var. , v. are taxonomic categories below the species level. In ,their use as species In level. below the categories taxonomic are = (e.g.: =variety complex et al. saxatilis 2012). They are considered infrasubspe areconsidered They 2012). (Fabricius, 1780) complex). 1780)complex). (Fabricius, = from the Latin rphological terms (Winston 1999). (Winston terms rphological stetit Caretta caretta but explicitly excluding subordinate taxathat may have been morph f. elongata " (but also "

with would Amphipoda include all the remaining complexus Mayr (1940) anddefined as "morphologically similar frasubspecific rank; see ICZN 1999: Art. 45.6), but 45.6),but Art. 1999: ICZN see rank; frasubspecific var. var. aberratio fic categories according to the ICZN (1999: Art. Art. (1999: ICZN tothe according categories fic Dautzenberg & P. , 1912). 1912). P.Fisher, & Dautzenberg olivacea olivacea form , form of ( ab. "), followed byanidentifier, is used. Deraniyagala, 1930). 1930). Deraniyagala, = aberration, abe Forma complector cific categories also in zoology zoology also in categories cific and varietas : "I entwine, encircle, encircle, : "Ientwine, rrant form).Both are the most most the are sensu 2014). Rosenbaum, secundum sp. inq.). This article isprotected Allrights bycopyright. reserved. Acceptedspecies inquirenda Article = in the sense used by (e.g.: by used sense the =in "circumscription" as intended by a particular auth a particular by intended as "circumscription" tothe taxon relates ataxonname ".taxon is as"taxon It known constitutes what a A view of ( aggregate" categories supraspecific informal qualifier the of use the to similarly complex, species the to reference a with ON in left be the and involved taxa the clarify to critical revision In practice, aspecies isverycomplex agroupof Pfenninger &Schwenk 2007). incr remarkably has complexes ofspecies number characters in individuals previously thought tobelong to the same species (Bickford is species ofcryptic existence The 1986). Knowlton emphasis on genetic relationship and the latte the and relationship genetic on emphasis put former the since subtle, is terms two the between The taxonomy. morphological Greek the (from species" been applied tocases of incomplete reproductive is in the literature. the in qualifier similar The ). ( name of species with unresolved taxonomicalstatus. Itcan be associated tothe term when attributing aspecimento acertain taxon.Instead, it is added byspecialist taxonomists tothe Glossary). Itcannot be considered anONqualifier,since it is notused tocommunicate uncertainty the id ofdoubtful aspecies indicates qualifier This inc. sed. inc. , abbreviation: ), which indicates that the taxonomic position taxonomic the that indicates ), which , abbreviation: sp. agg. sp.

sec. sec. or = according to, following (e.g.: (e.g.: following to, = according agg. sp. inq. Sousa ) and "species) and group" ( krypto species dubia =species tobe queried (e.g.: : "hidden") since they cannot be discriminated through classical comparable to species complex are used, including "species "species including used, are complex species to comparable (Osbeck, 1765) sensu Jefferson & Rosenbaum, 2014). 2014). Rosenbaum, & Jefferson sensu 1765) (Osbeck, ("doubtful species", abbreviated group or (Kennedy, Kukla & Paterson 2005; Franz, Peet & Peet& Franz, & 2005; Kukla or (Kennedy, Paterson r on morphological rese r on morphological entity, needing further investigation (ICZN 1999: 1999: (ICZN investigation further needing entity, generally inferred by diverging non-morphological non-morphological diverging by inferred generally diagnostic traits. In this case the identification may identification Inthiscase traits. may the diagnostic eased due to molecular tomolecular due eased similar, closely related sp related closely similar, olation. "Cryptic species" literally means "hidden "hidden means literally species" "Cryptic olation. Sousa chinensis , sp. gr. of the taxon is also uncertain (ICZN 1999: 1999: (ICZN isalsouncertain taxon ofthe Polycirrus pellucida Polycirrus or gr. (Osbeck, 1765) sec. Jefferson & & Jefferson sec. 1765) (Osbeck, ). mblance (Steyskal 1972; 1972; (Steyskal mblance studies (Knowlton 2000; 2000; (Knowlton studies ecies still waiting fora waiting still ecies sp. dub. (Quatrefages, 1866) 1866) (Quatrefages, et al. ) is also found incertae sedis incertae stetit 2007). The 2007).The . Other

This article isprotected Allrights bycopyright. reserved. Accepted Article interchangeably with an by tothe ofaname usage generally more refers term ofwhen taxon", "potential concept Anyway, equivalent introducingthe taxonconcept. the tothe the "sec." sign specified be may Taxonomic 2008). Weakley ( this approach is still debated, particularly when when particularly isstilldebated, approach this widening (see eg. Lepage, Vaidya & Guralnick 2014; Laurenne 2014;Laurenne Guralnick & Vaidya eg.Lepage, (see widening practice tospecify the taxon concept, e.g. good a be may it case, this In taxon. same the of circumscriptions different are there when meaningful definition explicit an with associated be therefore retraced onthe basis of published details. Inprinciple, taxonreported each in ataxonomic listshould with different names. Problems arise when the taxonconcept is notexplicitly stated, norit can be namemore new taxa taxon concept and rank,with with same simultaneous or of the the creation one comesfrom the partitiona taxon,which of causes theestablishmentmore circumscribed, anew, of case A typical views. authors or sources different to according exist simultaneously can concepts taxon different or evolve, can name a to associated concept taxon The process. identification taxonomic concepts related are not toON,they fundamental information about the Despite achecklist. amonograph key, article, or aresearch anidentification reference canbe the Accordingly, 2005). Paterson & Kukla (Kennedy, taxa other with relationships exclusive mutually to stated are which specimens) (or taxa subordinate taxon concept is implied (e.g. (e.g. isimplied concept taxon the to reference complete the when used also are superlatives, and comparatives related well as The "taxon concept" first arosefor at botanical databases, butits use in biodiversity informatics is Taxa atanyrankare circumscribedeitherse bya s.l. , s. lat. secundum , s.a.

, and a referenceand a Berendsohn authorship. tothe (1995) recommended this approach was traditionally used since the early 1800s (Welter-Schultes 2012). The term term The 2012). (Welter-Schultes 1800s early the since used traditionally was s. ampl. secundum ; "inthe; broadsense") and . Qualifiers with a looser meaning such as as such meaning alooser with . Qualifiers s.l., s.l., Cliona byreferring identification tothe key. s.s.). s.s.). applied to zoological data (Welter-Schultes, 2012). 2012). (Welter-Schultes, tozoologicaldata applied of the taxonconcept. Inpractice, this reference is author (ICZN, 1999: Glossary), but it is often used it used is often but author 1999:Glossary), (ICZN, sensu stricto t of diagnostic characters, the diagnostic bylisting set t of of belong tothe taxa or, finally, by asserting the by adding after the name a label composed of composed alabel name afterthe adding by et al. ( s.s. 2014; Franz 2014;Franz , s. str. sensu lato ; "in the strict sense"), as as sense"), strict the "in ; et al. 2015). However, However, 2015). or sensu amplo sensu

females (generally abbreviated abbreviated (generally females ovigerous of presence the as such used, are history life to related qualifiers other Sometimes recorded. For certain taxonomic groups, particularstages, lifesuch as This article isprotected Allrights bycopyright. reserved. abbreviated ( cast as such remains of occurrence the record may list taxonomic casesexual of dimorphisms, byapplying the habitual gender qualifiers ( adultus juvenile /ae can be diagnostic of environmental conditions. Main lifestage qualifiers include: stages life different at of presence the where , freshwater of biomonitoring the in instance Life stages also (see determinable objectively (abbreviated " term the case, this in alive; still were they when collected individuals same taxonomical(and ecological) meaning ofsingleparts detached body during sampling andhandling from ( Acceptedvalva/ae Article / / i , abbreviation: strictly necessary, it can be dropped toavoid redundancy. The qualifiercan differentiate be used to stages life particular with ecologicalsignificance. Ifnot are notfully developed (e.g.: both can be used asONqualifier account incomplete for to identificationwhen diagnostic characters of allow aproperinterpretation Both these qualifiers are used todifferentiate life stages particularwith ecological significance andto s , abbreviation: ), etc., sometimes), etc., with indications aboutthe degr , abbreviation: may be indicated byproper qualifiers.They frequently are used in certainareas of research, for ex. frag. ), empty mollusc shells ( shells mollusc empty ), or fr. lv. ) is usedON qualifier,) as indicatingthat the may quantifiable notbe and/or ad. juv. or (e.g.: (e.g.: (e.g.: (e.g.: l. (e.g.: ov. elegans , species morhua ovig. salinarius Chironomus ). The of the individuals is also sometimes recorded, particularly in particularly recorded, sometimes isalso individuals ofthe sex The ). the abundances, which may bevery theabundances, whichmay

conchylium/a indeterminabilis juv.). Linnaeus, 1758 juv.). 1758juv.). Linnaeus,

(Verrill, 1873) ad.). 1873)ad.). (Verrill, , abbreviated ). Kieffer, 1915 lv.). lv.). 1915 Kieffer,

/ae ee of conservation. These remains have notthe ( conch. p. ) in holometaboulous insects, may be be may insects, holometaboulous ) in , or" m., high but ephemeral. Moreover, Moreover, highbutephemeral. fragment ♂ conch ; f., ", " ", ♀ " or " or ). Moreover, some some ). Moreover, shell fragmentum/a ") andvalves exuvia/ae ,

This article isprotected Allrights bycopyright. reserved. taxonomistscan focus potential on non-indigenous previously or undescribedspecies, andset aside Kaiser 50%(e.g. than more even sometimes species, may be improved with anaccurate use of ON,as faunal listsmay include several unidentified oruncertain number of species still undescribed orpoorly known. In these cases, the quality of biodiversity assessments such assea, deep the the Antarcti the or still larg is biodiversity world's ofthe knowledge The 2009). Klump & (Huber degree confidence of in datacould be quantified, for instance byassigning them avalue in the [0,1] interval (Zingone quality ofinformation, traceability allow the uncertainty of the identification. Sta thing. ONqualifiersshould be as univocal andunambiguous as possible, since avague use increases the same the mean to used been have qualifiers different time, same the At inconsistently. applied been have different kinds anddegrees uncertainty. of Historically, ONqualifiers have been given multiple meanings, or notation ormay be included definitively in taxonomic listsmatrices. andSA This allows tool manage to identificationmeans by ofqualifiersnames. taxon combined ONqualifiers to beused asaprovisional can Open Nomenclature toolthatallows asemantic is scientists tocommunicate the of uncertainty ataxonomic Discussion Accepted timea when particular in efforts, methods, management andcost.A good time ofuncertainty during laboratory workbetter can direct research a quality The 2013). Vogler & (Chesters GenBank as such databases in deposited are sequences nucleotide whose specimens affects also identification intrinsically characterized higher by uncertaintyapplied, may be visual such census. as Uncertainty in the state ofpreservation. Whenthe sampling of organismswanted, is notpossible or other approaches requested toidentify alargenumber of specimens, including atdifferentstages life andin different technician including ofexpertise, levels different collect is data ofbiodiversity amount considerable A high degree uncertainty of in taxonomy is notlimited Article

et al. 2015). Moreover, since some ON terms may refl termsmay ON some since 2015).Moreover, nd resources are limited (Stribling, (Stribling, limited are resources nd ndardization of ON,together with the awareness past of practices, would facilitating the reappraisal of past ofpast reappraisal the facilitating nd resolution of taxonomic data is necessarily constrained by sampling sampling by constrained necessarily is data taxonomic of resolution nd s and parataxonomists (Janzen 2004; Krell 2004), are are 2004), Krell 2004; (Janzen parataxonomists s and et al. ed during biomonitoring programs. Staff members with Staffmembers biomonitoring programs. ed during ely incomplete (Costello, May & 2013). Ecosystems Stork & 2013).Ecosystems May (Costello, incomplete ely c region remain unexplored to a great extent, with a a with extent, great a to unexplored remain region c 2009; Baker 2009;Baker tothe frontiers of research. biodiversity Today, a et al. ect different levels of uncertainties, the the ofuncertainties, levels ect different records and the assessment of their oftheir assessment the and records Moulton & Lester 2003). Specialist Specialist 2003). Lester & Moulton 2012; 2012;Melo et al. 2014). 2014). software platforms, toextract, manage andintegrate biodiversity information (Berendsohn is aneed for dataandmetadata standards, controlled vocabularies andother semantic tools, as well as concepts, highly-structured ontologies are required (Laurenne Collaboration (Federhen 2012). Due toinherent complexity of taxonomic, nomenclatural andsystematic Taxonomy database provides anomenclatural frameworkto the International Nucleotide Sequence Database NCBI the instance for study; of fields ofother development tothe instrumental are them of Some 2013). databasesspecimens, for taxaand taxon names are in continuous development (Parr literatur legacy from data biodiversity of retrieval 2015).Au Krishtalka & Soberón Peterson, This article isprotected Allrights bycopyright. reserved. the Global Names Usage which Bank, is alist all publishedstatements aboutlifeforms,and the Global Names Pattersonindex content (http://globalnames.org/; aim is toprovide asingleshared platformcross-link to distributed biological data,using taxonomic names to framework(www.obofoundry.org).initiative Another Global is the(GNA), Architecture whose Names (Walls Ontology developed ontologies, suchas the Darwin-SW (Baskauff &Webb in press) and the Biological Collections Qualifier Standards (TDWG). They bothallow tostore ON Wieczorek berlin.de/TDWG/CODATA/default.htm; Holetschek and exchange of primary biodiversity datainclude the ABCD (http://bgbm3.bgbm.fu- observatio and i.e.specimens data, biodiversity primary ON signs donotform partof taxonnames(ICZN however 1999), they are partof core the information of required (see e.g. David al. points in time increasingly are digitalized andmade available through online(Hardisty resources Due tothe rapid development of biodiversity informatics,data from different sources, geographical and 2015). Occhipinti-Ambrogi Galil& (Marchini, species non-indigenous ON in left Taxa samples. preserved orpoorly incomplete

Accepted greater sametimes, 2012).Atthe Article "), aswell identification other as This totreat remarks.way ONqualifiers has been inherited bymore et al. 2012), both ratified and maintained inte bythe bothratified and 2012), et al. 2014),the latter being partof the Open Bi et al. 2012). 2012). and harmonization between biological disciplines is is disciplines biological between harmonization and communication tomated text mining and semantic semantic and textmining tomated e (Gerner, Nenadic & Bergman 2011; Lyal 2016). Large scale scale Large 2016). Lyal 2011; Bergman & Nenadic e (Gerner, et al. et al. qualifiers (as values of the specific term" specific ofthe (asvalues qualifiers ns (Chapman 2005). Main standards for the access to access forthe standards Main 2005). (Chapman ns are considered a critical point when compiling lists of of lists compiling when critical point a considered are 2012) and the Darwin Core (http://rs.tdwg.org/dwc/; Core(http://rs.tdwg.org/dwc/; Darwin the 2012)and 2010; Pyle 2016). Main components of the GNA are are GNA ofthe components Main 2016). Pyle 2010; et al. rnational organization Biodiversity Information Information Biodiversity organization rnational ological and Biomedical Ontologiesological andBiomedical Foundry 2014;Walls annotation of content foster the fosterthe ofcontent annotation et al. et al. 2014). Therefore, there there 2014).Therefore, 2012; Costello Costello 2012; et al. 2011;Parr Identification et al. 2013; 2013; et al. et

assistance with Latin . Finally, we thank wethank Finally, etymologies. Latin with assistance we Also suggestions. valuable forproviding Italy), Venice, Palma (UniversityBournemouth, ofMenegon UK), Ceregato,S. GuarneriDr.(CNR-ISMAR, Dr. andDr. A. I. It Padua, of (University Minelli A. Prof. thank sincerely We Acknowledgements non-taxonomists involved in biomonitoring andbiodiversitystudies. to the harmonization of ONterminology andpractices andtoprovide asynthetic guide for taxonomists and automated processing. However, this is beyond the scopeof this paper, whose main purpose is tocontribute This article isprotected Allrights bycopyright. reserved. AcceptedFlagship Project (SP3_WP2_AZ1_UO03) RITMARE MIUR. funded by that greatly improved the quality ofmanuscript. thegratefully We acknowledgesupport the from National the fo Explicit community. scientific the by agreed widely qualifiersconsistent onthe in todevelop legacy other, a dataand, vocabulary andsemanticON, for rules and integration of primary biodiversity data,onthe one handit is necessary toidentify the meaning ON of management the optimize To species. new a of establishment the with resolved later be may "aff." or "sp." as Articledoesn't take into account the dynamic nature of ON: for instance, the uncertainty expressed byqualifierssuch valuab of aloss in result may it time same atthe but aggregation ofdataatasafetaxonomic level.Thisoverestimating avoids biodiversity (Vandepitte et al. (Franz concepts taxonomic of different treatment the including issues, major some presents performed, automatically or manually either categories, evaluation (Zingone and databases, they need avery careful critical revision, with the aim of internal consistency andquality inventories taxonomic or time in integrated are sources different from data biodiversity primary When standardized, community-developed vocabulary of ONterms. the indexing and reconciling allow components GNA Other all vari of collection isa which Index, 2010). With respect to ON qualifiers, usually data-cle usually qualifiers, toON respect 2010).With

et al. 2015). The transfer of information from information of transfer The 2015). scientific of names, including surrogates andnames of taxaleft in ON. & Peet 2009), errors,synonyms spelling Peet2009), & the anonymous referees for rmalization would also ensure also would rmalization le taxonomic (and ecological) information. Moreover, it Moreover, information. ecological) (and taxonomic le aly), Dr. C. Glasby (MAGNT, Darwin, Australia), Dr. P. Darwin, Australia), (MAGNT, Dr. C.Glasby aly), aning procedures require just their removal and the the and removal their just require procedures aning would like to thanks A. Bracco (Genova, Italy) for content. All these tools lack a clear reference toa aclearreference toolslack Allthese content.

primary data to abstract taxonomic taxonomic toabstract data primary and taxa left in ON (Vandepitte (Vandepitte ON taxaleftin and their constructivetheir remarks data interoperability and and interoperability data et al. 2010) 2010)

This article isprotected Allrights bycopyright. reserved. () andrelated databases. coordination andstandardisation in marine biodiversity through the World Register of Marine Species Costello,Boxshall,Bouchet, M.J.,Fauchald, P., G., K.,Gordon,D.,Hoeksema, B.W. sequence datafor the construction of phylogenetic supermatrices. ambiguity Vogler,A.P.(2013)Resolving D.& Chesters, version 1.0 A.D.(2005) Chapman, No Zoological Y.(2015) Cambefort, species awindow as ondiversity andconservation. Ng, N.S., P.K.L.,Meier, D.J.,Sodhi, D.,Lohman, Bickford, Web -Interoperab Semantic Baskauf, S.J. &Webb, C.O. (in press) Darwin Core-based terms for expressing biodiversity dataas RDF. Series Progress Marine Distributional patterns of deep-seacoral assemblages in threesubmarine canyons off Newfoundland, . R. Haedrich, P.V.R., Snelgrove, V.E., Wareham, Baker, K.D., References Thismanuscript does notinclude anydata. Accessibility Data Accepted st From platforms: information Berendsohn,Güntsch, A.,Hoffmann, W.G., Kohlbecker,A. (2011) Biodiversity Müller, N., K. A.,Luther, & Berendsohn,(1995) W.G. Theconcept of "potentialdatabases. taxa" in Article Nomenclature. Open (1988) P. Bengtson, (eds K. Chemla & J. Virbel), pp. 143-181. 143-181. pp. J.Virbel), & (eds K.Chemla . Report for the Information Facility, Copenhagen. Copenhagen. Facility, Information Biodiversity Global forthe . Report

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Systematic Zoology durch Erläuterung der Internationalen Regeln rine biodiversity research: creating a a creating research: biodiversity rine et al. Journal of PLoS ONE (2014) Semanticssupport of in , 21,446. , 7(1), e29715. , 7(1), e29715. . Columbia University University Columbia . Hydrobiologia Journal of The ZooKeys , 644,1- , 550, , 7, , This article isprotected Allrights bycopyright. reserved. Accepted from the topdown. increases essentially confidence of degree The qualifiers. Nomenclature Open main using for Fig.1. Flowchart Article Appendix S1. Summary of Open Nomenclature qualifiers other main and qualifiers in alphabetical order. Additional Supporting Information may be found in the online version of this article. Supporting Information Methods in Ecology and Evolution

Open Nomenclature in the biodiversity era

Marco Sigovini, Erica Keppel and Davide Tagliapietra

Appendix S1. Summary of Open Nomenclature qualifiers and other main qualifiers in alphabetical order. Table 1 . Summary of Open Nomenclature (ON) qualifiers. Main abbreviations ("signs") found in literature are reported. Definitions follow main literature as well as recommendations presented in this paper.

Term Sign Meaning and notes Examples affinis , aff. "Has affinity with". The specimen has some Pourtalesia aff. alcocki Koehler, 1914 species affinis sp. aff. affinity to a known species but it is not identical Petrolisthes sp. nov. aff. rufescens to it; it generally implies distinction more than a (Heller, 1861) possible identity, in contrast with the qualifier confer ; see also species proxima and ex grege . It is often used in combination with the ON qualifier species nova . Also used in the sense of confer (this usage is discouraged, for details see the main text ). animalia cetera a.c. It groups all the unidentified specimens that are a.c. A.C. not listed as separate taxa. The term cetera Peracarida c. (abbreviated c. or cet. ) may be applied to a given high-rank taxon, meaning that identification at a lower taxonomic level has not been attempted (see also stetit ) but explicitly not including subordinate taxa that may have been identified. confer cf. "Compare with". Specimens should be compared Polycera cf. hedgpethi Er. Marcus, 1964 cfr. to reference material, since most of the conf. diagnostic characters correspond to a given sp. cf. species but some are unclear. Also used in the sense of affinis and species incerta (these usages are discouraged, for details see the main text ). ex grege ex gr. "Of the group including". The specimen has Pseudocandona ex gr. eremita gr. some affinity to a known species or it belongs to (Vejdovsky, 1882) a species group or species complex; see also affinis and species proxima . familia genus species fam. gen. sp. The specimen has not been attributed to any Nematoda fam. gen. sp. (= Nematoda sp.) known species nor family; see also species . genus et species nova gen. et sp. nov. The specimen is considered to belong to a new Vulcanoctopus hydrotermalis gen. nov., sp. nov. species and a new genus; for more details, see gen. et sp. nov. nov. gen. et sp. species nova . etc. genus novum gen. nov. The specimen is considered to belong to a new, gen. nov. novum genus g. nov. previously undescribed, genus; for more details, gen. n. see species nova . g. n. nov. gen. etc. genus species gen. sp. The specimen has not been related to any Zoarcidae gen. sp. (= Zoarcidae sp.) g. sp. known species nor genus; see also species . species sp. The specimen has not been identified, nor it has Nucula sp. been related to any known species; the sp. A uncertainty is potentially provisional: it could be due to the lack of suitable dichotomous keys, or to the occurrence of a species not previously described. Also used in the sense of species indeterminabilis and stetit (these usages are discouraged, for details see the main text ). species (pl.), spp. More than one species belonging to the same Unio spp. species plurimae sp. pl. genus (or higher-rank taxon) are included. species incerta ? The identification is uncertain; it usually Tubifex tubifex (Müller, 1774) ? sp. inc. indicates a higher reliability with respect to = Tubifex tubifex (Müller, 1774) sp. inc. confer . The sign "sp. inc." is also used in the sense of species , species indeterminabilis and species inquirenda (these usages are discouraged, for details see the main text ).

2 Table 1 . Summary of Open Nomenclature (ON) qualifiers. (cont.)

Term Sign Meaning and notes Examples species indeterminabilis indet. The specimen is indeterminable beyond a Lekanesphaera indet. (also reported as ind. certain taxonomic level due to the deterioration Clavulariidae indet. species indeterminata ) sp. indet. or lack of diagnostic characters. Also used in the sp. ind. sense of species and stetit (these usages are discouraged, for details see the main text ). species nova sp. nov. The specimen is considered to belong to a new, Pelagia benovici sp. nov. nova species spec. nov. previously undescribed species. (1) When Tisbe sp. nov . sp. n. describing a new species, the use of the qualifier nov. sp. is required by the ICZN (1999) to explicitly nov. spec. indicate the taxa name as intentionally new. n. sp. (2) Used as ON qualifier to refer to a new, still unnamed species before the formal publication of the description. species proxima prox. The specimen is near to a known species but it is prox. costata species near sp. prox. not identical to it; see also affinis and ex grege . Tiwari & Biswas, 1952 nr. sp. nr. stetit stet. Identification at a lower taxonomic level has not Teredinidae stet. = Teredinidae or the intentional been attempted, even if allowed by the sample absence of qualifiers conditions. It may also be used when more records with different ON qualifiers need to be merged at a safe taxonomic level. subspecies ssp. The only infraspecific rank regulated by the ICZN Salmo trutta Linnaeus, 1758 ssp. subsp. (1999). As ON qualifier, it indicates that the specimen probably belongs to a subspecies but it has not been related to any known one; see also species .

3 Table 2 . Summary of other main qualifiers cited in the paper. Main abbreviations ("signs") found in literature are reported. Definitions follow main literature as well as recommendations presented in this paper.

Term Sign Meaning and notes Examples aberratio ab. Aberration, aberrant or atypical form, which deviates in discretus ab. woerndlei important characters from the . Zimmermann, 1919 Infrasubspecific category according to the ICZN (1999); since infrasubspecific categories are not regulated by the ICZN, it should be regarded as a morphological term; see also morph. adultus /i ad. Life stage. (Verrill, 1873) ad. forma f. Form. Infrasubspecific category according to the ICZN (1999) f. elongata (subspecific in case of names published before 1961 and not Dautzenberg & P. Fisher, 1912 expressly given infrasubspecific rank by the author); since infrasubspecific categories are not regulated by the ICZN, it should be regarded as a morphological term; see also varietas . fragment, frag. Detached body part. The organism may not be objectively frag. fragmentum/a fr. determinable and quantifiable; see also species indeterminabilis (Table 1). incertae sedis inc. sed. A taxon of uncertain taxonomic position (ICZN 1999); the Polychaeta inc. sed. qualifier is often attached to a high-rank taxon to include all the species in the group with uncertain taxonomic position. juvenile /s juv. (1) Life stage. (2) Used as ON qualifier when diagnostic Gadus morhua Linnaeus, 1758 juv. characters necessary to identification are not fully developed. Acartia juv. larva /ae lv. (1) Life stage. (2) Used as ON qualifier when diagnostic Chironomus salinarius Kieffer, 1915 lv. l. characters necessary to identification are not fully developed. lv. morph morph (1) Specimen within a species differing in consequence of cavernosa (Linnaeus, 1767) form intrapopulational polymorphism. (2) Infrasubspecific category morph 1 according to the ICZN (1999); since infrasubspecific categories are not regulated by the ICZN, it should be regarded as a morphological term; see also aberratio . secundum sec. "According to". It is followed by a reference to the taxon Sousa chinensis (Osbeck, 1765) circumscription such as an , a checklist or a sec. Jefferson & Rosenbaum, 2014 research article; it is a label for the applied taxon concept. sensu sensu "In the sense used by". It refers to the usage of a name in a Sousa chinensis (Osbeck, 1765) sense different from that of other authors (ICZN, 1999); it is sensu Jefferson & Rosenbaum, 2014 generally used in the same sense of the qualifier secundum . Cliona s.l., Cliona s.s. Qualifiers with a looser meaning include sensu latu ( s.l. , s. lat. ) or sensu amplo ( s.a. , s. ampl. ) and sensu strictu ( s.s. , s. str. ), together with related comparatives and superlatives. species complex complex (1) A group of related species characterized by unclear (Fabricius, 1780) boundaries, typically due to phenotypical similarities (sibling or complex cryptic species). (2) A group of very similar, closely related taxa still waiting for a critical revision. Also used as ON qualifier in a sense similar to stetit (Table 1). Other informal supraspecific categories include species aggregate ( sp. agg. or agg. ) and species group ( group , sp. gr. or gr. ). species dubia sp. dub. A species of doubtful identity, needing further investigation; the Maera aequimana Ledoyer, 1979 sp. dub. taxonomic status of the species is unresolved; see also species inquirenda . species inquirenda sp. inq. A species of doubtful identity, needing further investigation Polycirrus pellucida (Quatrefages, 1866) (ICZN, 1999); the taxonomic status of the species is unresolved; sp. inq. see also species dubia . varietas var. Variety. Infrasubspecific category according to the ICZN (1999) Caretta caretta var. olivacea v. (subspecific in case of names published before 1961 and not Deraniyagala, 1930 expressly given infrasubspecific rank by the author); since infrasubspecific categories are not regulated by the ICZN, it should be regarded as a morphological term; see also forma .

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