Introduction to Biosystematics - Zool 575

Introduction to Biosystematics Outline Lecture 5 - Nomenclature 1. Naming

2. History

3. Nomenclature vs

4. Rules - ICZN

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Biosystematics Biosystematics - readings

Lecture 5: Nomenclature & Classification

Mayr, E.& P. D. Ashlock (1991) Principles of Systematic Zoology, 2nd Edition, McGraw-Hill, Inc., NY. pp. 383-406. character identification evolution Smith, H. M. and O. Williams (1970) The salient provisions of the International Code of Zoological Nomenclature: A summary for nontaxonomists. Bioscience, descriptions phylogeny 20: 553-557

**Winston, J. E. (1999) Describing : Practical Taxonomic Procedure for Biologists. Columbia University Press, NY. pp. 19-40, 407-432. collections classification biogeography

Describing species = assigning names to groups (populations) = classification 3 4

Naming Naming

Basically a two step process Step 1 - “Is it a new species?” (minimum)

1. Taxonomist finds something thought to 1. Are the diagnostic (unique) characters constant be unnamed, “new” across large samples within the ?

2. Have you compared the new species with 2. A name is given by publishing descriptions of all its congeners? (globally?) (according to the rules of nomenclature) 3. If the group has not been revised (well), have you at which point the name is introduced to the examined the primary specimens of literature all congeners?

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Naming Naming

Step 1 - “Is it a new species?” (better) Comment on uniqueness…

1. Do you have samples large enough to obtain - Early taxonomists dealt mostly with the statistically significant differences in quantitative “obvious” cases - wide phenotypic gaps traits? - Current taxonomists deal more & more with 2. Do you have DNA data that indicates a “gap” exists between the new species and congeners? difficult cases: cryptic species, incipient species, etc. - narrow gaps 3. Do you have data that indicate reproductive barriers exist? (e.g. courtship songs, pheromones) - Taxon dependent (e.g. birds vs insects)

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Naming History of the Code

Step 2 - Publishing Linnaean system of binominal nomenclature

- See lecture 9 for format of paper - Vast improvement over phrase names and prior naming systems BUT… - Rules of International Code of Zoological Nomenclature must be followed (for - No stability, Linnaeus & users of his system ) - this lecture would change names to “improve” them (e.g. change the name to better reflect distribution)

- Lamarck, 1798, criticized lack of rules, instability, & chaos under the Linnaean 9 system 10

History of the Code History of the Code

Problem: no rules to ensure species were Priority - oldest name is used known by a single name - Starting at a fixed date: - Early 1800s exploration of tropics revealed - 1758 - zoology (except spiders in Clerck’s immense & surprising diversity 1757 publication) - 1753 - botany - To stabilize names, in 1813 a Swiss botanist, Augustin Pyramus de Candolle, - Exceptions exist for cases when principle introduced the concept of Priority of priority would result in extreme, if temporary, instability

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History of the Code ICZN - goals for the code

The first Zoological Code Conflicting goals of taxonomic classification (not nomenclature) - 1843 the “Strickland” code - provide a unique, stable name - Formed by members of the Strickland (ideally, would never change) Commission, including Charles Darwin - provide a natural classification - Revisions resulted in two competing codes (requires constant change as new data reveal new relationships) - A governing body, International Commission for Zoological Nomenclature, 1895 13 14

ICZN - goals for the code Key elements of the ICZN 1. Promote stability (International Code of Zoological Nomenclature) 2. Promote universality 1. Neutrality - doesn’t infringe on taxonomic 3. Names will be unique & distinct judgement - By establishing rules for: 2. Will not solve rank placement problems - Publication - Priority Nomenclature ! Taxonomy - Typification

Current Code: edition 4, 1999. 15 16

More Key elements of the ICZN Nomenclature: Provisions for the formation (International Code of Zoological Nomenclature) and use of a system of names (rules) 1. Typification - the name-bearing type, all e.g. nomenclatural status of a name - its names are tied to a type specimen standing in nomenclature, does it conform to the rules? 2. - oldest name is valid Taxonomy: The theory & practice of 3. Principle of Stability - case by case basis to classifying organisms (opinions) prefer stability over priority (in rare cases) e.g. taxonomic status of a name - is it valid, ICZN regulates names from superfamily to is it unique to one species? 17 subpsecies 18

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Priority & Stability Priority & Stability

Priority works to 1) Recognize first scientist to publish and Nomina oblita (a nomen oblitum) - “forgotten name” - many cases of unknown publications 2) Promote stability because there can only be one “first” publication whereas there an older name not used in over 50 years can be many arguments for “better” can be suppressed as a nomen oblitum if names following the principle of priority would destabilize usage 3) However, there are exceptions… imagine if someone found an older name for Drosophila melanogaster! 19 20

Priority & Stability ICZN Rules

Nomina oblita (a nomen oblitum) - “forgotten Failure to follow the rules of proper name” publishing can result in a nomen nudum - a “naked name,” a name that was not e.g. Nicrophorus americanus Olivier 1790 published properly - fails to be established - well known name a nomen nudum is not available Nicrophorus orientalis Herbst 1784 - never used name an has been published properly, is available for use as a taxon When invoked the becomes a name nomen protectum 21 22

ICZN Rules - publication ICZN Rules - publication

To be available a name must: To be available a name must:

1. Be published in the meaning of article 8 2. Be spelled using only the 26 letters of the 8.1.1 + 8.1.2 + Latin alphabet “8.1.3 it must have been produced in an edition containing simultaneously obtainable copies by a method 3. Be a word (e.g. not ‘cbafdg’) that assures numerous identical and durable copies” Also recommendations (not requirements): Web pages do not assure identical or e.g. do not use unmodified vernacular durable copies (common) names, or offensive names

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ICZN Rules - Validity Example of simple synonymy list (the kind you will be creating) Valid name - the single correct, accepted Nicrophorus americanus Olivier, 1790 name for a taxon Synonyms: Silpha (Nicrophorus) orientalis Herbst,1784:77 [nomen oblitum] Many names might be available for a species, Nicrophorus americanus Olivier, 1790:(no. 10):6 but there is only one valid name Nicrophorus virginicus Frölich, 1792:123 Necrophorus grandis Fabricius, 1792a:247 [type: ZMUC, 2 specimens] If there are multiple names for one species Valid name = senior these are synonyms of each other other, invalid synonyms = junior synonyms

*Taxonomy tells us which names are Some species, e.g. swan mussel, synonyms; nomenclature tells us which cygnea, have hundreds of synonyms (400+) of the synonyms is the valid name* 25 26

Example of bibliographic synonymy list Reading a Synonomy list (table)

Nicrophorus insularis Grouvelle, 1893: 161 Type locality: “Sumatra”, Indonesia Methia necydalea (Fabricius) NEW COMBINATION Type depository: MNHN: Paris [!] 1893 Grouvelle: 161 (JA: descriptions in French) >> Orig. comb.: Necrophorus insularis [!] 1903 Portevin: 331 (JA: descriptions in French) >> as valid species 1920 Portevin: 399 (JA: descriptions in French) >> as valid species Saperda necydalea Fabricius, 1798: 148 1922 Portevin: 55 (JA: descriptions in French) >> New syn./status as N. nepalensis var. insularis 1923 Portevin: 307 (JA: key, descriptions in French) >> as N. nepalensis var. insularis 1926a Portevin: 208 (Book: revision: key, descriptions & catalog in French) >> as N. nepalensis var. insularis 1928 Hatch: 129 (Book Chapter: catalog in English) >> as N. nepalensis var. insularis 1964b Hlisnikovsky: 244 (JA: descriptions in German) >> New status as N. nepalensis form insularis [ab. intended?] Saperda necydalina; Fabricius, 1801: 332. Schoenherr, 1975 Emetz & Schawaller: 230 (JA: checklist in German (English summary)) >> as N. nepalensis ab. insularis 1990 Hanski & Niemelä: 149 (Book Chapter: in English) >> New status as valid species 1817:439 UNJUSTIFIED EMENDATION 1991 Hanski & Krikken: 195 (Book Chapter: in English) >> as valid species 2001 Peck: 94 (JA: in English) >> as valid species herein >> desig. lectotype = humeralis Pic, 1917: 2 Methia pusilla; Salle, 1889: 468 (not Newman, 1840). Type locality: “Java” [-from label] Type depository: MNHN: Paris [!] MISIDENTIFICATION 1917a Pic: 2 (JA: descrip. of var. in French) >> Orig. comb.: Necrophorus insularis var. humeralis; auth. attrib. Port. [!] 1920 Portevin: 399 (JA: descriptions in French) >> descrip. of variation: 1 unavailable name 1926a Portevin: 253 (Book: revision: key, descriptions & catalog in French) >> New syn. of N. nepalensis var. insularis 1928 Hatch: 129 (Book Chapter: catalog in English) >> as syn. of N. nepalensis var. insularis 1993 Kozminykh: 67 (JA: key, descriptions, regional in Russian) >> as syn. of N. nepalensis Thia jamaicensis Gahan 1902: 44. NEW SYNONYMY herein >> REVISED SYNONYM of N. insularis Grouvelle

Objective synonyms = same type specimen

Subjective synonyms = different type specimens 27 28

Homonyms Homonyms Two or more available names having the secondary homonymy - later combined same spelling established for different taxa with the same generic name

primary homonymy - originally identical Ptomascopus morio Kraatz 1887 Careospina snail genus Nicrophorus morio Motschulsky 1841 Careospina moth genus OK.. Until P. morio is moved into the genus secondary homonymy - later combined Nicrophorus by Chapman in 1955 with the same generic name Then: N. morio (Kraatz, 1887) - homonym homonyms across the plant & of N. morio Motschulsky 1841 kingdoms are OK eg. Pieris & Pieris 29 30

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26 Homonyms in Nicrophorus Homonyms Without the author, a binomen is not necessarily unique!

N. basalis Faldermann, 1835 N. marginatus Gistel, 1857 junior homonym - younger of two N. basalis Gistel, 1848 N. marginatus Fabricius, 1801 homonyms N. bipunctatus Kraatz, 1880 N. maritimus Guérin-Ménéville, 1835 N. bipunctatus Portevin, 1914 N. maritimus Mannerheim, 1843

N. cadaverinus Gravenhorst, 1807 N. orientalis Motschulsky, 1860 senior homonym - oldest of two + N. cadaverinus Gistel, 1857 N. orientalis (Herbst,1784) N. cadaverinus Mareuse, 1840 homonyms N. plagiatus (Ménétries, 1854) N. insularis Grouvelle, 1893 N. plagiatus Motschulsky, 1870 N. insularis Lafer, 1989 N. morio (Kraatz, 1887) - note parentheses added N. pollinctor LeConte, 1854 N. interruptus Brullé, 1832 N. pollinctor Mannerheim, 1853 N. interruptus Gistel, 1857 to author name when species is no longer in N. interruptus Stephens, 1830 N. quadricollis Hatch, 1928 original genus N. quadricollis Gistel, 1848 N. lunatus Fischer von Waldheim, 1842 N. lunatus LeConte, 1853 In Botany the author who moves the name gets appended also e.g. Rosa alba (L.) Fab. 31 32

Types example

“name bearers” - anchors to all formal names Dr. Fraunhofer has representative for the population been studying a snail species, Regardless of opinions on species Hendersonia demarcations / limits / boundaries the type occulata. always belongs to the name with which it was published She finds this species has been cited 37 A new name must have a declared type times in the specimen (article 72.3 ICZN ed 4) as of 1999 literature.

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example example Dr. Fraunhofer decides Further study reveals to check the type that the species specimen & finds it in everyone thought to the Linnaean be H. occulata isn’t collection in London The actual H. occulata Surprisingly, the turns out to be rare & looks like unstudied and the this species everyone thought to be H. occulata is 35 undescribed! 36

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Why types? Why not descriptions? Types Descriptions are interpretations of observations Primary types - single specimen linked to name Types are things (objective) 1. Holotype - one specimen chosen by author in original description

2. Lectotype - one specimen taken from a type series during revisionary work

3. Neotype - all types lost, reviser may designate a new specimen as primary No type & poor description = nomen dubium37 type 38

Types - secondary types Types - locality Many older names have no holotype but instead have a type series = The locality from which the holotype was collected is the type locality when a lectotype is chosen from a series the remaining specimens become This is the best place to collect specimens if paralectotypes one is designating a neotype

when a holotype is designated, other Note on priority - if two names have same date specimens used become & month is unknown, “first reviser” gets to choose the valid name These are secondary types and have no ‘legal’ standing 39 40

Types above species group Rank Taxa Suffix *Kingdom Animalia The type of a genus is the *Phylum Arthropoda *Class Insecta *Order Diptera Superfamily -oidea *Family Culicidae -idea - anchors the genus name Subfamily -inae Tribe -ini a) original designation Subtribe -ina b) subsequent designation *Genus Culex c) monotypy (only one species in genus) *Species Culex pipiens

Type of the family is the type genus 41 * Mandatory ranks 42

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Classification written works concerning animals published unpublished Femoria (Ectodermia) populella Available unavailable suppressed Genus (subgenus) specific epithet NAMES NOMENCLATURAL ACTS

Scientific vernacular VALID invalid suppressed (=Brontosaurus) giganteus Available Unavailable Excluded Genus (synonym of genus) specific epithet Potentially valid objectively invalid

Jr objective synonym Jr homonym suppressed Informal groupings: species group subjectively invalid species complex VALID Jr subjective synonym nomina dubia jr homonym suppressed

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Terms - from lecture & readings You should be able to

Priority Describe the conflicting goals of the code of nomenclature Stability Primary type ICZN Holotype Describe the difference between nomenclature and taxonomy Typification Lectotype Nomen oblitum Neotype Describe the difference between an available name and a Nomen nudum Secondary type valid name Available (name) Valid (name) Paralectotype Understand a synonymy table Synonyms Syntype Basically, describe any of the terms on the previous slide (senior/junior, Type locality objective, subjective) Type species Explain why type specimens are so important Primary & secondary Type genus homonymy 45 46

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