BioConcepts

subspeciesSubspezies (ger.)

• A population, or group of populations, that occupies a geographically closed subdivision of the range of the species and that consists of individuals with distinct characters, with at least 75% of the individuals being separable, on the basis of the particular character, from all (99+ per cent) of the members of the other populations of the species.
  category species see also race

  1780

  Scheinarten, Halbarten oder Subspecies

  Ehrhart, F. (1780). Versuch eines Verzeichnisses der um Hannover wildwachsenden Pflanzen. Hannoverisches Magazin 18, 209-240: 211.

  1781

  Subspezies (Untergattungen, Raçes)

  Esper, E.J.C. (1781). De varietatibus specierum in naturae productis. Sectio I.: 19 (§XIV).

  1784

  Halbarten, Scheinarten, Subspecies. [...] Sie sind, mit einem Wort Varietates constantes, oder ein Mittel zwischen Arten und Spielarten. Sie unterscheiden sich von Arten, daß sie in kleinen und weniger beträchtlichen Umständen voneinander abgehen; und von Spielarten differiren sie, daß sie sich beständig durch den Saamen fortpflanzen, und immer wieder ihres gleichen zeugen

  Ehrhart, F. (1784). Botanische Bemerkungen. Hannoverisches Magazin 22, 113-128; 129-144; 161-176: 169.

  1866

  Fast bei allen Gruppen von Organismen haben sich […] die besseren und gewissenhafteren Systematiker genöthigt gesehen, von denjenigen Arten, die genauer bekannt und in sehr zahlreichen Exemplaren untersucht sind, und namentlich von denjenigen, welche einen sehr grossen Verbreitungsbezirk besitzen, die abweichenderen Individuen, welche die specifischen Charaktere mehr oder weniger modificirt zeigen, oder sich als mehr oder minder entschiedene Uebergangsbildungen zu verwandten Arten hinneigen, als besondere Unterarten (Subspecies) oder Spielarten (Varietätes) zu beschreiben

  Haeckel, E. (1866). Generelle Morphologie der Organismen, vol. II, 337-8.

  1891

  Subspecies sind Formen, welche von anderen noch nicht genügend entfernt sind, um ihnen den Rang der Species beizumessen. […] Ich glaube […], dass man gut thun wird, Formen, welche sich nur durch geringe Grössenunterschiede, hellere oder dunklere Farbentöne, solche kleine Unterschiede in der Zeichnung, welche leicht variabel sind, wie z. B. ein etwas grösserer Fleck einer Farbe hier und dort (sofern sie überhaupt ein bestimmtes Gebiet bewohnen und in diesem beständig sind!), als Subspecies zu betrachten, auch wenn man die intermediären Formen nicht vor sich hat.

  Hartert, E. (1891). Katalog der Vogelsammlung im Museum der Senckenbergischen Naturforschenden Gesellschaft in Frankfurt am Main: xiii; xiv.

  1903

  With subspecies we designate [...] nothing else but the geographically separated different components of one and the same type, which components represent together a species. The criterion of a subspecies is not a certain amount of difference, but bodily difference and geographical separation. Synoecic varieties—i.e. varieties from the same locality—are never subspecies.

  Rothschild, W. & Jordan, K. (1903). A revision of the lepidopterous family Sphingidae: xlii.
  

  1903

  Mit Subspezies bezeichnen wir die geographisch getrennten Formen eines und desselben Typus, die zusammengenommen eine Spezies ausmachen. Es ist also nicht etwa ein geringes Maß an Unterschieden, das uns bestimmen darf, eine Form als Spezies aufzufassen, sondern Unterschiede verbunden mit geographischer Trennung, natürlich bei allgemeiner Uebereinstimmung

  Hartert, E. (1903). Einleitung. In: Die Vögel der paläarktischen Fauna, vol. 1, 1. Lieferung, I-XI: VI.

  1935

  If […] those subspecies are considered invalid that cannot be identified in at least 75% of the specimens, then a prediction [of the number of species] is possible

  Mayr, E. (1935). How many birds are known? Proceedings of the Linnaean Society of New York 45-46, 19-23: 23.

  1942

  In the case of subspecies, it is a good convention that at least 75 percent of the individuals of one subspecies (or of the available specimens) should be separable, on the basis of their diagnostic characters, from the specimens of the most similar subspecies

  Mayr, E. (1942). Systematics and the Origin of Species: 16.

  1949

  the 75 per cent rule […] is here defined to mean that 75 per cent of a population must be separable from all (99+ per cent) of the members of overlapping populations to qualify as a subspecies. An equivalent statement is that 97 per cent of one of two overlapping populations must be separable from 97 per cent of the other

  Amadon, D. (1949). The seventy-five percent rule for subspecies. Condor 51, 250-258: 258.

  1953

  the subspecies concept is the most critical and disorderly area of modern systematic theory – more so than taxonomists have realized or theorists have admitted. Particular confusion surrounds the drawing of the lower limits of the subspecies category within that spectrum of classes recognized by Mayr as extending from “the local population into the subspecies.” The difficulties in this delimitation stem from four outstanding features of geographical variation: (1) the tendency for genetically independent characters to show independent geographical variation; (2) the capacity for characters to recur in more than one geographical area, yielding polytopic races; (3) the common occurrence of the microgeographical race; (4) the necessary arbitrariness of any degree of population divergence chosen as the lowest formal racial level

  Wilson, E.O. & Brown, W.L. Jr. (1953). The subspecies concept and its taxonomic application. Systematic Zoology 2, 97-111: 100.

  1963

  A subspecies is an aggregate of local populations of a species, inhabiting a geographic subdivision of the range of the species, and differing taxonomically from other populations of the species

  Mayr, E. (1963). Animal Species and Evolution: 348.

  1976

  Subspecies, like all taxonomic categories except the species, are subjective

  Amadon, D. & Short, L.L. (1976). Treatment of subspecies approaching species status. Syst. Zool. 25, 161-167: 164.

  1982

  The thousands of formally named subspecies of birds range in distinctiveness from groups of populations barely discernible on the basis of weak divergence in a single character to geographic forms that illustrate trenchant differences in morphology, coloration, and voice. With such a breadth of variation covered by one taxonomic umbrella, small wonder that biologists continue to question the meaning and usefulness of this category. The heart of the problem lies in the fact that the complexity of natural variation below the species level defies easy organization; it is a mixed bag. Furthermore, each species seems to show a unique pattern

  Johnson, N.K. (1982). Retain subspecies – At least for the time being. The Auk 99, 605-606: 605.

  1982

  subspecific names should not be used to describe populations differing only through smooth clines reflecting general primary intergradation. […] Subspecies […] should be used in two situations: (1) allopatric populations where definition of the populations is clear, distinct, and total (or very nearly so); and (2) situations where secondary contact between distinct populations has occurred and the zone of intergradation is relatively narrow

  Monroe, B.L., Jr. (1982). A modern concept of the subspecies. The Auk 99, 608-609: 609.

  1982

  subspecies 1: A group of interbreeding natural populations differing taxonomically and with respect to gene pool characteristics, and often isolated geographically, from other such groups within a biological species q. v., and interbreeding successfully with these groups where their ranges overlap. 2: A taxon at the rank of subspecies.

  Lincoln, R.J., Boxshall, G.A. & Clark, P.F. (1982). A Dictionary of Ecology, Evolution and Systematics: 238.

  1982

  it seems curious that qualitative examination of color or a few skin measurements of a few specimens, often without statistical tests for clines or without adequate sampling of intermediate geographical areas, frequently results in trinomials, while the authors of large, quantitative studies frequently avoid them. This strongly suggests to me that most subspecies are not to be taken too seriously

  Barrowclough, G.F. (1982). Geographic variation, predictiveness, and subspecies. The Auk 99, 601-603: 601.

  1982

  the subspecies fulfilled a most important historical role by undermining the essentialistic species concept and also by contributing to a far better understanding of the geographic variation of species taxa in nature. […] Today, however, the primary use of subspecies is as a sorting device in collections, that is as an index to populations that differ from each other “taxonomically”

  Mayr, E. (1982). Of what use are subspecies? Auk 99, 593-595: 595.

  1992

  Subspecies. Within a species, a named, recognized allopatric subpopulation which is (still) genetically compatible with other subpopulations, but is set apart by a concordant array of genetic and phenotypic characters

  Amadon, D. & Short, L.L. (1992). Taxonomy of lower categories – suggested guidelines. In: Monk, J.F. (ed.). Avian Systematics and Taxonomy (= Bulletin of the British Ornithologists’ Club 112A), 11-38: 12.

  2002

  In essence, a subspecies is a collection of populations occupying a distinct breeding range and diagnosably distinct from other such populations […]. [T]o be a valid subspecies 75% of a population effectively must lie outside 99% of the range of other populations for a given defining character or set of characters

  Patten, M.A. & Unitt, P. (2002). Diagnosability versus mean differences of sage sparrow subspecies. The Auk 119, 26-35: 27.

  2004

  “subspecies” warranting formal recognition could […] be conceptualized as groups of actually or potentially interbreeding populations (normally mostly allopatric) that are genealogically highly distinctive from, but reproductively compatible with, other such groups. Importantly, the empirical evidence for genealogical distinction must come, in principle, from concordant genetic partitions across multiple, independent, genetically based molecular (or phenotypic; Wilson and Brown 1953) traits

  Avise, J.C. (1994/2004). Molecular Markers, Natural History, and Evolution: 362-3.

  2004

  Mitochondrial DNA sequence data reveal that 97% of continentally distributed avian subspecies lack the population genetic structure indicative of a distinct evolutionary unit. […] [I]t is illogical that, at the subspecies level, formal taxonomic names can be applied to trivial non-historical patterns of variation. Only taxa defined by the congruence of multiple morphological or molecular characters should be recognized at some rank. Over 90% of continental avian subspecies fail this test. Thus, avian taxonomists must revise classifications by eliminating thousands of subspecies names

  Zink, R.M. (2004). The role of subspecies in obscuring avian biological diversity and misleading conservation policy. Proceedings of the Royal Society of London, Series B 271, 561-564: 561; 563.

  2010

  I regard the concept [viz. the subspecies] as a useful convenience. However, I submit that art and judgment will always be involved in practice and that no one trinomial treatment can be scientifically proved to be the biologically correct one. In this context, the subspecies concept itself is simply too heterogeneous to be classified as strict science

  Fitzpatrick, J.W. (2010). Subspecies are for convenience. Ornithological Monographs 67, 54-61: 54.

  2010

  From personal experience in attempting to use subspecies diagnoses, such as the keys in the Birds of North and Middle America series (Ridgway and Friedmann 1901-1950), I predict that more than 75% of North American subspecies taxa delimited by mensural data would not survive application of the 75% rule [...]. A subspecies is a distinct population, or group of populations, that occupies a different breeding range from other populations of the same species; individuals are distinguishable from those other populations by one or more phenotypic traits at the 95% level of diagnosability

  Remsen, J.V. (2010). Subspecies as a meaningful taxonomic rank in avian classification. Ornithological Monographs 67, 62-78: 64f.