Introduction to Biosystematics - Zool 575 Introduction to Biosystematics Outline Lecture 5 - Nomenclature 1. Naming 2. History 3. Nomenclature vs Taxonomy 4. Rules - ICZN 1 2 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 species: 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 genus? 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 type specimens of literature all congeners? 5 6 1 Introduction to Biosystematics - Zool 575 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) 7 8 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 animals) - 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 11 12 2 Introduction to Biosystematics - Zool 575 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. Principle of Priority - 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 3 Introduction to Biosystematics - Zool 575 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 available name has been published properly, is available for use as a taxon When invoked the valid name 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 23 24 4 Introduction to Biosystematics - Zool 575 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 synonym these are synonyms of each other other, invalid synonyms = junior synonyms *Taxonomy tells us which names are Some species, e.g. swan mussel, Anodonta 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