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The Classification of Avian Species and Subspecies

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The Classification of Avian Species and Subspecies The classification of avian species and subspecies A lucid presentation of the baseson which speciesare lumped or split John T. Ratti INTRODUCTION (1971:99) reported that ornithologists evidencethat these two indigo birds have "universallyadopted" the biologi- locally behave as distinct biological ECENTLY,THERE HAVE BEENa cal speciesconcept, defined by Mayr species"(Payne 1973:175). number of changesin the classifica- (1969:20) as "groups of interbreeding tion of bird speciesapproved by the natural populations that are reproduc- ISOLATING MECHANISMS Committee on Classification and No- tively isolatedfrom other suchgroups." menclature (American Ornithologists' Strictly and simply, this definition states N THEPARAGRAPH above I referred Union Check-List Committee 1973, that if two populationsoccur in the same to "isolating mechanisms." This con- 1976). Nearly all changesinvolve two or region (sympatric) and do not inter- cept should be discussedbefore con- more closelyrelated species or subpopu- breed, they are separate species sidering evolution of species.Popula- lations of a species.Thus, the systema- regardlessof phenotypic(appearance), tions of birds that occur in the same tist rarely has difficulty separating morphological, or ecological similari- generalregion, yet fail to interbreed,are speciesfrom different orders or families, ties. On the other hand, if two sympatric said to have isolating mechanisms-- and challengesin proper classification or adjacentpopulations frequently inter- "propertiesof individualswhich prevent usually involve populations of the same breed, they are the same species(con- interbreeding" (Mayr 1970:56). Mayr genus or species.Such changesare cas- specific)regardless of phenotypic, mor- (1970) classifiedisolating mechanisms as ually referred to as "lumping" (combin- phological, or ecological differences. "premating" or "postmating." Premat- ing two or more speciesinto a singlespe- Consequently, "reproductive isolation" ing isolating mechanisms include cies) or "splitting" (separating a species is the key to the biological speciescon- seasonaland habitat isolation, behavior- into two or more species) and often cept. However, this does not mean that al isolation, and mechanical isolation. generate confusion and misunderstand- closely related valid species never Postmatingmechanisms include gametic ing among amateur ornithologists and hybridize. Isolating mechanisms are mortality, zygotic mortality, hybrid in- some professional biologists. To often imperfect, and Mayr (1951:102) viability, and hybrid sterility. Avlan ehmlnate this confusion one must study noted that "I understand the occasional isolating mechanisms are commonly and considerpast and present concepts interbreeding of two otherwise well- reported to be behavioral premating of speciation.The following brief review delimitedsympatric species. There is no mechanisms associated with species- may be helpful in this regard. Some conceptual difficulty in regard to this specificrecognition of song, color pat- readersmay wish to review the glossary type of hybridization." Speciationis re- terns, courtship displays, and similar of technicalterms in the Appendix. gardedas essentiallycomplete if, during mechanismsthat may be reinforced by sympatry, interbreedingis reducedto a conditioningor imprintingby newlyhat- THE SPECIES CONCEPT level that preventsgenetic swamping by ched chicks (Beach and James 1954, the parent species(Mayr 1959, Bigelow Marler 1957 and 1961, Hinde 1959, Im- OSTSPECIES OF BIRDScurrently 1965).An examplemay be helpful at this melmann 1975a and 1975b). recognized were described on a point. Because two speciesof indigo Smith (1966) found contrastingcolor morphological basis, i.e., classification birdsof Africa, Vidua chalybeataand V. patternsnear the eyeand wingtippattern of speciesaccording to consistent size, purpurascens, were morphologically in- to be the isolatingmechanisms among structure, and color differences. When distinguishable,their status as separate four speciesof gulls (Larus spp.). Jehl dealing with extinct species and fossil species was questioned (White' 1962, and Bond (1975) describeda similarsitu- records, we are forced to base classifica- Payne 1973). To clarify the problem, ation with murrelets of the genus En- tion on morphological characteristics. Payne (1973) studied the degreeof re- domychura and suggestedthat facial However, most modern systematistsdis- productive isolation; pair observations patterns near the eye and bill shapeboth favor the morphological speciesconcept revealed 71 chalybeata x chalybeata, 5 function as isolating mechanismsbe- becauseit may lead to incorrect classifi- chalybeatax purpurascens,and 73 pur- tween theseclosely related species.Song cation (of subspecies or failure to purascens x purpurascens. Although a and/or call notes have been identified as recognize the separate speciesstatus of small number of hybrids was observed, isolatingmechanisms for syrupatticmea- slbhngspecies). These problemswill be thesedata, along with phenotypicdiffer- dowlarks (Sturnella spp.), Australian discussedlater with examples. Selander ences,were reported to provide "direct Wedgebills(Psophodes spp.), flycatch- 860 AmericanB•rds, November 1980 ers (Empidonax spp.), and indigo birds occasionallyinterbreed. If hybridization the Galapagos Islands and Hawaiian of the sub-genusHypochera (Szijj 1966, is frequent, the populationswill quickly Honey Creepers(Drepanididae) are dra- Ford and Parker 1973, Stein 1963, and lose any differences evolved during iso- maticexamples of founderspecies. Mayr Traylor 1966, respectively).Combina- lation, and their speciesstatus will be (1951:9•)suggested that foundersmay tions of song and color pattern have unchanged.However, if isolatingmech- speciateby double invasion. Where "an been identified as isolating mechanisms anisms prevent interbreeding, or only island is repeatedlyinvaded by colonists by Gill and Murray (1972) and Brown allow for occasionalhybridization, then from a distant mainland, it may happen (1967) while Payne (1973) reportedthe a new specieswill have evolved (Dobz- that the descendants of the first wave of combination of song and behavior. hansky 1937, Lack 1944, Miller 1947, colonists have changed so much that Selanderand Giller's (1961:77) study Mayr 1951 and 1970, Sibley 1961, they are reproductively isolated from of the Great-tailed (Quiscalus mex- Selander 1971, Stebbins 1971, and Bush new arrivals." tcanus)and Boat-tailed grackles(Q. ma- 1975). Speciationis usually a long-term jor) reported the primary isolating processand likely involvesthousands of SUBSPECIES mechanismto be female recognition of years of evolution. Figure 1 was con- male "behavioral differences, both at structed to offer a graphic review of ECAUSESPECIATION issuch a long- term process,some populations can- the time of nest site selection and at time geographicspeciation. not be classifiedas simply belongingto of mating." Many other isolatingmech- At this point one shouldcarefully con- one speciesor another. Certain popula- anisms are reported, including habitat sider the following note of caution. tions are in intermediatestages of specia- (Brewer 1963) and seasonal isolation Divergenceand developmentof isolating tion or have specialmorphological char- (Smith 1966). mechanisms are not mutually depen- acteristics(Mayr 1951, 1963, and 1970, An important part of isolatingmecha- dent; many populations are known that Selander 1971, Stebbins 1971). msms and speciation is reinforcement have diverged morphologically in allo- The taxonomiccategory of subspecies (Mayr 1970). For example, a population patty but possessno isolating mechan- is assignedto distinct subpopulationsof having color patterns as an isolating isms in zones of sympatry. These popu- the same species.This category ranges mechanism may have a fractional por- lations are usually called subspecies. from populations having barely percep- tion of members that will readily Conversely, some populations evolve ef- tible morphologicaldifferences to those hybridize. However, if the hybrid off- fective isolating mechanisms, become that appear sufficiently distinctive to spring(Fl) expressan intermediatecolor sympatric, but fail to diverge mor- have been classifiedas separatespecies. pattern, they may be selectedagainst by phologically or in general appearance. In each case, the populations so classi- the majority in their attempt to find a Such populations are called sibling fied have discrete geographic ranges, mate. This negative selection will even- species. have morphologically diverged, but have tually minimize or completelyeliminate Mayr (1970:278) noted that, given the not developed sufficient isolating hybridization. overwhelmingacceptance of geographic mechanisms to maintain reproductive speciation, "the basic problem of THE SPECIATION PROCESS isolation should their ranges overlap. speciation consists in explaining the Consequently, most bird populations AYR(1970:247-277) discussed various origin of the gaps between sympatric that undergo geographicspeciation like- modesof speciationwith examples species."With the exceptionof founder ly evolve through a period when they from several taxonomic groups (birds, species(to be discussed),extrinsic bar- would be recognized as subspecies plants, insects, etc.), but ornithologists riers promoting speciation result from (Figure 1). Hence, we do not expect to agreethat geographicdivergence best ex- large-scale environmental change, such observe subspeciesthat are sympatric
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