Systematics - BIO 615

Home , Nomen dubium, Swan mussel, Syntype, Valid name (zoology)

Systematics Lecture 5 - Nomenclature Names are tags, like codes or passwords, that form a link between nature and our information about nature.

If this link is broken information is lost.

“Nomina si nescis, perit et cognito rerum”

[Who knows not the names, knows not the subject] Linnaeus, Criticia Botanica, 1773.

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Scathophaga impudicum

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Biosystematics

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 7 8

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, i.e. it is ‘official’ NOTE: currently the publication can be in any of a large variety of formats all congeners? and is not required to be registered with a central database of names 9 10

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 Winston (1999) Describing species - 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 13 system 14

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 change if new data reveal new relationships) - A governing body, International Commission for Zoological Nomenclature, 1895 17 18

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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 judgment - By establishing rules for: 2. Will not solve rank placement problems - Publication - Priority Nomenclature ≠ Taxonomy - Typification

Current Code: edition 4, 1999. 19 20

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? 21 subspecies 22

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! 23 24

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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 25 26

ICZN Rules - publication ICZN Rules - publication

Amendment of Articles 8, 9, 10, 21 and 78 - ZooKeys 219: 1– To be available a name must: 10, doi: 10.3897/zookeys.219.3944

1. Be published in the meaning of article 8 “The requirements for electronic publications are that:

8.1.1 + 8.1.2 + 1) The work be registered in ZooBank before it is published “8.1.3 it must have been produced in an edition containing simultaneously obtainable copies by a method 2) That the work itself state the date of publication and contain that assures numerous identical and durable copies” evidence that registration has occurred

3) That the ZooBank registration state both the name of an Web pages do not assure identical or electronic archive intended to preserve the work and the durable copies (but see Amendment of Articles 8, 9, 10, 21 and 78 - ISSN or ISBN associated with the work. ZooKeys 219: 1–10, doi: 10.3897/zookeys.219.3944) 27 28

ICZN Rules - publication ICZN Rules - publication Recent case of the new monkey To be available a name must: described without a type specimen - photo only. 2. Be spelled using only the 26 letters of the Latin alphabet Jones et al. 2005. The Highland Mangabey Lophocebus kipunji: a new species of African monkey. Science. 3. Be a word (e.g. not ‘cbafdg’) Caused a debate - Justified due to conservation status? Also recommendations (not requirements): Small population size? e.g. do not use unmodified vernacular (common) names, or offensive names Bad idea to describe new species based on photos only. 29 30 No voucher for independent verification.

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ICZN Rules - publication ICZN Rules - Validity Valid name - the single correct, accepted name for a taxon

“It’s no good to science if it’s Many names might be available for a species, running around free.” but there is only one valid name

Link Olson If there are multiple names for one species these are synonyms of each other

*Taxonomy tells us which names are synonyms; nomenclature tells us which 31 of the synonyms is the valid name* 32

Example of simple synonymy list Example of bibliographic synonymy list

Nicrophorus insularis Grouvelle, 1893: 161 Nicrophorus americanus Olivier, 1790 Type locality: “Sumatra”, Indonesia 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 Synonyms: 1920 Portevin: 399 (JA: descriptions in French) >> as valid species 1922 Portevin: 55 (JA: descriptions in French) >> New syn./status as N. nepalensis var. insularis Silpha (Nicrophorus) orientalis Herbst,1784:77 [nomen oblitum] 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 Nicrophorus americanus Olivier, 1790:(no. 10):6 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?] Nicrophorus virginicus Frölich, 1792:123 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 Necrophorus grandis Fabricius, 1792a:247 [type: ZMUC, 2 specimens] 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 Valid name = senior synonym Type locality: “Java” [-from label] Type depository: MNHN: Paris [!] 1917a Pic: 2 (JA: descrip. of var. in French) >> Orig. comb.: Necrophorus insularis var. humeralis; auth. attrib. Port. [!] other, invalid synonyms = junior synonyms 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 Some species, e.g. swan mussel, Anodonta herein >> REVISED SYNONYM of N. insularis Grouvelle cygnea, have hundreds of synonyms (400+) Objective synonyms = same type specimen

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Reading a Synonomy list (table) Homonyms Two or more available names having the Methia necydalea (Fabricius) NEW COMBINATION same spelling established for different taxa Saperda necydalea Fabricius, 1798: 148 primary homonymy - originally identical Saperda necydalina; Fabricius, 1801: 332. Schoenherr, 1817:439 UNJUSTIFIED EMENDATION Careospina snail genus Careospina moth genus Methia pusilla; Salle, 1889: 468 (not Newman, 1840). MISIDENTIFICATION secondary homonymy - later combined Thia jamaicensis Gahan 1902: 44. NEW SYNONYMY with the same generic name

homonyms across the plant & animal 35 kingdoms are OK eg. Pieris & Pieris 36

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

secondary homonymy - later combined N. basalis Faldermann, 1835 N. marginatus Gistel, 1857 with the same generic name N. basalis Gistel, 1848 N. marginatus Fabricius, 1801 N. bipunctatus Kraatz, 1880 N. maritimus Guérin-Ménéville, 1835 N. bipunctatus Portevin, 1914 N. maritimus Mannerheim, 1843

Ptomascopus morio Kraatz 1887 N. cadaverinus Gravenhorst, 1807 N. orientalis Motschulsky, 1860 N. cadaverinus Gistel, 1857 N. orientalis (Herbst,1784) Nicrophorus morio Motschulsky 1841 N. cadaverinus Mareuse, 1840 N. plagiatus (Ménétries, 1854) N. insularis Grouvelle, 1893 N. plagiatus Motschulsky, 1870 N. insularis Lafer, 1989 OK.. Until P. morio is moved into the genus N. pollinctor LeConte, 1854 N. interruptus Brullé, 1832 N. pollinctor Mannerheim, 1853 Nicrophorus by Chapman in 1955 N. interruptus Gistel, 1857 N. interruptus Stephens, 1830 N. quadricollis Hatch, 1928 N. quadricollis Gistel, 1848 N. lunatus Fischer von Waldheim, 1842 Then: N. morio (Kraatz, 1887) - homonym N. lunatus LeConte, 1853 of N. morio Motschulsky 1841 37 38

Without the genus or author, an epithet is not Homonyms necessarily unique! NOTE: epithet ≠ species name junior homonym - younger of two Among the ~7,000 species of non- homonyms marine Alaskan arthropods: senior homonym - oldest of two + alaska 2 The epithet qualiﬁes homonyms alaskae 6 the genus name alaskaensis 2 so using an epithet N. morio (Kraatz, 1887) - note parentheses added alaskana 8 alone is like saying to author name when species is no longer in alaskanum 2 ʻredʼ instead of saying original genus alaskanus 9 ʻred wagonʼ alaskense 4 In Botany the author who moves the name gets alaskensis 64 appended also e.g. Rosa alba (L.) Fab. 39 40

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. 41 42

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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 & holotype looks like unstudied and the this species everyone thought to be H. occulata is 43 undescribed! 44

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 dubium 45 type 46

Types - secondary types Types - locality Many older names have no holotype but instead have a type series = syntypes The locality from which the holotype was collected is the type locality when a lectotype is chosen from a syntype 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 paratypes & month is unknown, “first reviser” gets to choose the valid name These are secondary types and have no ‘legal’ standing 47 48

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Types above species group Rank Taxa Suffix *Kingdom Animalia The type of a genus is the *Phylum Arthropoda *Class Insecta *Order Diptera type species Superfamily -oidea *Family Culicidae -idae - 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

(note that species has two names) Type of the family is the type genus 49 * Mandatory ranks 50

The family group name when writing in English Classification is "a noun in the nominative plural" Femoria (Ectodermia) populella Genus (subgenus) specific epithet Examples: "The Tenebrionidae are widespread." Apatosaurus (=Brontosaurus) giganteus "The tenebrionids are widespread." Genus (synonym of genus) specific epithet "That tenebriond is black.”

NEVER "That Tenebrionidae is black.” Informal groupings: species group (see also the word ‘data’) species complex

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written works concerning animals Terms - from lecture & readings published unpublished

Available unavailable suppressed Priority Nomen dubium

NAMES NOMENCLATURAL ACTS Stability Primary type ICZN Holotype Scientific vernacular VALID invalid suppressed Typification Lectotype Available Unavailable Excluded Nomen oblitum Neotype Potentially valid objectively invalid Nomen nudum Secondary type Jr objective synonym Jr homonym suppressed Available (name) Paratype Valid (name) Paralectotype subjectively invalid Synonyms (senior/ Syntype Type locality VALID Jr subjective synonym nomina dubia jr homonym suppressed junior, objective, subjective) Type species Primary & secondary Type genus 53 homonymy 54

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You should be able to

Describe the conflicting goals of the code of nomenclature

Describe the difference between nomenclature and taxonomy

Describe the difference between an available name and a valid name

Understand a synonymy table

Explain any of the terms on the previous slide

Explain why type specimens are so important

Know how many names are in a species name 55