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The Taxonomic Rank of Spizella Taverneri: a Response to Mayr and Johnson Ornithology Program (HRC) Harry Reid Center for Environmental Studies 5-2001 The taxonomic rank of Spizella taverneri: A response to Mayr and Johnson John Klicka University of Nevada, Las Vegas, [email protected] Robert M. Zink University of Minnesota Jon C. Barlow Royal Ontario Museum W. Bruce McGillivray Provincial Museum of Alberta Terry J. Doyle Ten Thousand Islands National Wildlife Refuge Follow this and additional works at: https://digitalscholarship.unlv.edu/hrc_ornithology Part of the Genomics Commons, Ornithology Commons, and the Population Biology Commons Repository Citation Klicka, J., Zink, R. M., Barlow, J. C., McGillivray, W. B., Doyle, T. J. (2001). The taxonomic rank of Spizella taverneri: A response to Mayr and Johnson. Condor, 103(2), 420-422. Available at: https://digitalscholarship.unlv.edu/hrc_ornithology/32 This Letter to the Editor is protected by copyright and/or related rights. It has been brought to you by Digital Scholarship@UNLV with permission from the rights-holder(s). You are free to use this Letter to the Editor in any way that is permitted by the copyright and related rights legislation that applies to your use. For other uses you need to obtain permission from the rights-holder(s) directly, unless additional rights are indicated by a Creative Commons license in the record and/or on the work itself. This Letter to the Editor has been accepted for inclusion in Ornithology Program (HRC) by an authorized administrator of Digital Scholarship@UNLV. For more information, please contact [email protected]. 420 COMMENTARY The Condor 103:420±422 q The Cooper Ornithological Society 2001 lidad de la reproduccioÂn, vocalizaciones y morfologõÂa apoyaban el reconocimiento de S. taverneri como una especie tanto ®logeneÂtica como bioloÂgica. Nada en el comentario de Mayr y Johnson causa que cambiemos THE TAXONOMIC RANK OF SPIZELLA esa conclusioÂn. Creemos que es probable que ese par TAVERNERI: A RESPONSE TO MAYR AND de taxones alopaÂtricos esteÂn aislados. Contrariamente 1 a Mayr y Johnson, creemos que se proporciona maÂs JOHNSON informacioÂn designando a S. taverneri como especie, pues revela el hecho de que ambos son taxones que JOHN KLICKA,2 Marjorie Barrick Museum of Natural evolucionan independientemente. La clasi®cacioÂn de History, Box 454012, University of Nevada Las Ve- Spizella debe mostrar el estatus de especies hermanas gas, 4505 Maryland Parkway, Las Vegas, NV 98154- de S. taverneri y S. breweri, sin tomar en cuenta para 4012 el balance el grado de divergencia de las secuencias entre las especies, como fue sugerido por Mayr y John- ROBERT M. ZINK, J. F. Bell Museum of Natural His- tory, 100 Ecology Building, University of Minnesota, son. St. Paul, MN 55108 Mayr and Johnson (2001) do not dispute the evidence JON C. BARLOW, Royal Ontario Museum, 100 Queen's presented by Klicka et al. (1999) that corroborated the Park, Toronto, ON M5S 2C6, Canada existence of a taxon named taverneri. The discussion W. B RUCE MCGILLIVRAY, Provincial Museum of Alber- instead concerns whether this taxon should be ranked ta, 12856-102 Avenue, Edmonton, AB T5N 0M6, Can- as a species or a subspecies. Mayr and Johnson believe ada that our evidence is best interpreted to indicate that taverneri is a subspecies of the biological species S. TERRY J. DOYLE, Ten Thousand Islands National Wild- breweri. We concluded that taverneri represents a life Refuge, 3860 Tollgate Blvd., Suite 300, Naples, FL newly evolved phylogenetic species, and probably a 34114 biological species as well. We believe that the argu- ments presented by Mayr and Johnson, grounded in Abstract. Mayr and Johnson suggest that Spizella the school of evolutionary systematics (Mayr and Ash- taverneri should be a subspecies of the biological lock 1991), are equivocal, and show why the biologi- species S. breweri, because it is possibly not repro- cal species concept (BSC) has been losing favor in ductively isolated. We originally concluded that evi- many disciplines (Wheeler and Meier 2000), including dence from mitochondrial DNA sequences, habitat preferences, timing of breeding, vocalizations, and ornithology (Zink and Davis 1999). Below we respond morphology supported the recognition of S. taverneri to the issues raised by Mayr and Johnson. as a phylogenetic and biological species. Nothing in Evidence for reproductive isolation. Mayr and John- the commentary by Mayr and Johnson causes us to son state that if genetic independence cannot be deter- change that conclusion. We believe that it is probable mined by direct observation (i.e., a ``test of sympat- that these two allopatric taxa are isolated. Contrary ry''), it must be inferred. This, in fact, has long been to Mayr and Johnson, we believe that more infor- recognized as a critical ¯aw in the biological species mation is given by ranking S. taverneri as a species, concept: the main criterion for ranking taxa as species because it reveals the fact that they are independently must be inferred in the thousands of cases (such as the evolving taxa. The classi®cation of Spizella should present one) in which diagnosable populations or convey the sister-species status of S. taverneri and S. groups of populations are allopatric. The methods of breweri, without regard for balancing the degree of inference are vague and vary among taxonomists. To- sequence divergence among species, as suggested by day, analyses of DNA are used to discover genetically Mayr and Johnson. isolated groups of populations and to recover their phylogenetic relationships. In modern systematics, the Key words: Brewer's Sparrow, classi®cation, DNA phylogeny is used for classi®cation (GutieÂrrez et al. sequences, species concepts, Timberline Sparrow. 2000), including the assignment of species limits (Zink and Davis 1999). Furthermore, it is now known that El Estatus TaxonoÂmico de Spizella the pattern of reproductive compatibility is not neces- taverneri: una Respuesta a Mayr y sarily an accurate predictor of phylogenetic relation- Johnson ship because non-sister taxa often hybridize (Zink and McKitrick 1995, Klicka et al. in press). Thus, species Resumen: Mayr y Johnson sugieren que Spizella limits based on recovered patterns of evolutionary his- taverneri debe ser una subespecie de la especie bio- tory (e.g., phylogenetic analyses of DNA) can con¯ict loÂgica S. breweri, porque posiblemente no se encuentra with those based on patterns of reproductive compat- aislada reproductivamente. Nosotros originalmente ibility. Zink and Davis (1999) suggested that the ap- concluimos que la evidencia de las secuencias del propriate research program was to map patterns of re- ADN mitocondrial, preferencias de haÂbitat, tempora- productive compatibility and isolation onto a phylog- eny. In this way, one can understand which character- istics contribute to reproductive compatibility and 1 Received 10 January 2001. Accepted 12 January isolation among taxa, rather than to use these catego- 2001. ries as ambiguous characters to diagnose species. 2 E-mail: [email protected] Our DNA analyses suggested that S. breweri and S. COMMENTARY 421 taverneri are recently isolated sister taxa. Whether they taken by Mayr and Johnson would have more validity are reproductively isolated is of interest, but is only if all subspecies were as clearly de®ned as S. taverneri relevant to determining whether S. taverneri is a bio- (although this would still not be an argument in favor logical species. Mayr and Johnson speculate that there of keeping the BSC, Zink and Davis 1999). Unfortu- is insuf®cient evidence to predict whether pairings be- nately, all subspecies are not created equal. The liter- tween individual S. taverneri and S. breweri would ature is fraught with poorly de®ned and indistinguish- yield viable offspring, were they to encounter each able subspecies. A growing body of evidence indicates other during the breeding season. We think that some that as many as 50% of avian subspecies are not cor- of the differences that help diagnose S. taverneri are roborated by mtDNA data, presumably because they relevant to the question of reproductive compatibility. are based on arbitrary divisions of single morpholog- For example, Mayr and Johnson disregard substantial ical character clines (Zink et al. 2000). For example, differences in the timing of breeding. Spizella taver- Curve-billed Thrashers (Toxostoma curvirostre) have neri does not return to the breeding grounds until 4± been divided into two morphologically distinct sub- 6 weeks after S. breweri has begun nesting, reducing species clusters (the Curvirostre and Palmeri groups) the likelihood of interbreeding. Mayr and Johnson also that correspond to an east-west division in southwest- emphasized the lack of information on pairing behav- ern North America, yet seven subspecies are generally ior of the two taxa. In particular, they focused on our accepted. Recent morphometric (Rojas-Soto 1998) and discussion of the differing vocal characteristics. Al- mtDNA analyses (Zink and Blackwell-Rago 2000) though differences in advertising song might or might support division into only two (or possibly three) evo- not be important in assessing reproductive isolation lutionary units, which we consider species. Indeed, a among allopatric populations (McKitrick and Zink loss of information occurs when all 7 subspecies are 1988), the vocal differences are more convincing than given equal rank. Similarly inappropriate is the clas- Mayr and Johnson suggest. Spizella taverneri songs si®cation of S. taverneri as a subspecies.
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