Passerellidae

Baird’s sparrow Ammodramus bairdii, copyright Gerardo Marrón.

Belongs within: Emberizoidea.
Contains: Spizella, Amphispiza, Chlorospingus, Peucaea, Ammodramus, Arremonops, Arremon, Passerella, Junco, Zonotrichia, Atlapetes, Pipilo, Melozone, Aimophila, Passerculus, Melospiza.

Sparrows of the West
Published 10 August 2021

Recent decades have seen significant shifts in the classification of birds, particularly among the Passeriformes, the perching birds. These shifts have lead to the recognition of a number of major groups that were previously obscured. Among these recent elevations are the New World sparrows of the Passerellidae.

Gambel’s white-crowned sparrow Zonotrichia leucophrys gambeli, copyright Gregory Smith.

The New World sparrows are part of a broader radiation known as the nine-primaried songbirds, along with such luminaries as finches, tanagers, and their Old World namesakes. The name ‘nine-primaried’ refers to the number of well-developed primary feathers (the long outer ones) in the wings; most other perching birds have ten distinct primaries. Though the nine-primaried songbirds have long been recognised as a coherent group, there has been a lot of disagreement over their subdivision. Historically, these subdivisions were strongly influenced by different bill shapes representing different diet specialisations, but recent molecular phylogenies have demonstrated that bill shape is more labile than previously recognised. The New World sparrows were usually regarded previously as a subgroup of the generalist seed-eating family Emberizidae, along with the buntings of the Old World, but molecular phylogenies have asserted the division between the hemispheres. Not all New World representatives of the old Emberizidae have shifted to the Passerellidae: a significant component of the Neotropical fauna (including the finches of the Galapagos islands) have instead proven to be closer to the fruit-eating tanagers of the Thraupidae. As currently recognised, the passerellids are a fairly coherent group of about 140 species distributed around North and South America.

Golden-winged sparrow Arremon schlegeli schlegeli, copyright Nick Athanas.

In general, the passerellids are small birds with simple, conical bills. Most are dull brownish in coloration though many are strikingly patterned, particularly around the head. Some are more distinctive: the South American sparrows of the genus Arremon often stand out as particularly colourful. Most passerellids are fairly retiring in their usual habits, foraging at or close to ground level. As noted before, they are mostly generalist feeders. Their short bills are excellently suited for milling the small seeds which make up a large part of their diet. However, they will also take insects and other small invertebrates. One widespread North American species, Ammodramus savannarum, has earned the vernacular name of “grasshopper sparrow” as a result. Notable outliers dietwise are the Neotropical bush-tanagers of the genus Chlorospingus which are primarily berry feeders. These largely greenish birds were previously classified with the Thraupidae as a result before molecular data led to their reassignment.

Common bush-tanager Chlorospingus flavopectus, copyright Becky Matsubara.

Whereas Neotropical members of the Passerellidae are mostly sedentary, North American species are often migratory, moving north with the approach of summer. However, migration is commonly related to environmental conditions. A number of species are migratory in the northern parts of their range but may be found in their southern territories year-round. In some species, migrating populations will leap-frog over resident populations, moving further south than any resident individuals during the winter months. Many passerellid species are strong singers and courting males will often select an exposed branch to sing from in contrast to their usual skulking habits. Other species, particularly those inhabiting open habitats where trees and shrubs are in short supply, may have prominent aerial displays. Males of one of these latter species, the lark bunting Calamospiza melanocorys, moult during the breeding season into black plumage with contrasting white patches on the wings and tail. During the remainder of the year, they are dull in coloration like their females. Nesting is conducted close to ground level like feeding with the nest often being a small cup in the ground concealed under vegetation. Where breeding has been studied in detail, passerellids are commonly what has been called “socially monogamous”. Males and females will form what appear to be monogamous pairs with one male remaining close to one female (though construction of the nest and incubation are done by the female alone). However, genetic studies on nestlings have found that chicks are not uncommonly not the child of their apparent ‘father’, indicating that females have not remained faithful to their mate.

Yellow-striped brush-finch Atlapetes citrinellus, copyright Ron Knight.

Prior to molecular studies, authors had suggested a possible division of North American passerellids between two evolutionary lineages based on ecology and behaviour, the grassland and brushland sparrows. A molecular study of passerellids by Klicka et al. (2014) identified eight well-supported clades within the family. Two further species, the large-footed finch Pezopetes capitalis of Central America and the Zapata sparrow Torreornis inexpectata of Cuba, were not robustly assigned to a clade. Identified relationships were comparable to but not entirely congruent with prior hypotheses. For instance, most ‘brushland sparrows’ (of the genera Passerella, Zonotrichia and Junco) belonged to a single clade but the remaining ‘brushland’ genus Melospiza was placed in a clade mostly made up of ‘grassland’ species. The diverse South American genus Arremon was supported as monophyletic but others were not. In particular, the North American Ammodramus was divided between two widely separated clades. This lead to the resurrection of the genus Ammospiza for a group of saltmarsh-breeding species. Deeper relationships within the family deserve further investigation.

Systematics of Passerellidae
PasserellidaeOF19
|--+--SpizellaOF19
| `--+--AmphispizaBF16
| `--+--Chondestes Swainson 1827BF16, B94 [Chondestinae]
| | `--C. grammacus (Say 1822) [=Fringilla grammaca]RJ11
| | |--C. g. grammacusRJ11
| | `--C. g. strigatus Swainson 1827 [incl. C. g. quillini]RJ11
| `--Calamospiza Bonaparte 1838BF16, B94 [Calamospizinae]
| `--C. melanocorys Stejneger 1885RJ11
`--+--ChlorospingusBF16
`--+--+--+--PeucaeaBF16
| | `--AmmodramusOF19
| `--+--ArremonopsBF16
| `--Rhynchospiza Ridgway 1898BF16, RJ11
| |--R. stolzmanni (Taczanowski 1877) [=Haemophila stolzmanni, Aimophila stolzmanni]RJ11
| `--R. strigiceps (Gould 1839) [=Zonotrichia strigiceps, Aimophila strigiceps]RJ11
| |--R. s. strigicepsRJ11
| `--R. s. dabbenei (Hellmayr 1912)RJ11
`--+--ArremonBKB15
`--+--+--+--PasserellaOF19
| | `--Spizelloides arboreaBF16
| `--+--JuncoBKB15
| `--ZonotrichiaBKB15
`--+--+--+--Pezopetes Cabanis 1861BF16, RJ11
| | | `--P. capitalis Cabanis 1861 [=Atlapetes capitalis]RJ11
| | `--+--AtlapetesBKB15
| | `--PipiloBKB15
| `--+--MelozoneBKB15
| `--AimophilaBF16
`--+--+--Oriturus Bonaparte 1850BKB15, RJ11
| | `--O. superciliosus (Swainson 1838) [=Aimophila superciliosa]RJ11
| | |--O. s. superciliosusRJ11
| | `--O. s. palliatus (van Rossem 1938)RJ11
| |--Artemisiospiza Klicka & Banks 2011RJ11
| | |--A. belli (Cassin 1850)RJ11 [=Emberiza belliRJ11, Amphispiza belliBKB15]
| | | |--A. b. belliRJ11
| | | |--A. b. canescens (Grinnell 1905)RJ11
| | | |--A. b. cinerea (Townsend 1890)RJ11
| | | `--A. b. clementeae (Ridgway 1898)RJ11
| | `--A. nevadensis (Ridgway 1874) [=Poospiza belli nevadensis; incl. A. n. campicola]RJ11
| `--Pooecetes Baird 1858BF16, RJ11
| `--P. gramineus (Gmelin 1789) [=Fringilla graminea]RJ11
| |--P. g. gramineusRJ11
| |--P. g. affinis Miller 1888RJ11
| `--P. g. confinis Baird 1858 [incl. P. g. altus]RJ11
`--+--+--PasserculusBF16
| `--+--‘Ammodramus’ bairdii (Audubon 1844)BKB15, RJ11 [=Emberiza bairdiiRJ11]
| `--+--MelospizaBKB15
| `--Xenospiza Bangs 1931BKB15, RJ11
| `--X. baileyi Bangs 1931 [=Ammodramus baileyi, Emberiza baileyi]RJ11
`--+--‘Ammodramus’ henslowii (Audubon 1829)BKB15, RJ11 [=Emberiza henslowiiRJ11]
| |--A. h. henslowiiRJ11
| |--A. h. houstonensisRJ11
| `--A. h. susurrans (Brewster 1918)RJ11
`--+--‘Ammodramus’ leconteii (Audubon 1844)BF16, RJ11 [=Emberiza leconteiiRJ11]
`--+--‘Ammodramus’ maritimus (Wilson 1811)BKB15, RJ11 [=Fringilla maritimaRJ11]
| |--A. m. maritimusRJ11
| |--A. m. fisheri Chapman 1899RJ11
| |--A. m. juncicola (Griscom & Nichols 1920)RJ11
| |--A. m. macgillivraii (Audubon 1834)RJ11
| |--A. m. mirabilis (Howell 1919)RJ11
| |--A. m. nigrescensOM91
| |--A. m. pelonotusLG91
| |--A. m. peninsulae Allen 1888RJ11
| `--A. m. sennetti Allen 1888RJ11
`--+--‘Ammodramus’ caudacutus (Gmelin 1788)BKB15, RJ11 [=Oriolus caudacutusRJ11]
| |--A. c. caudacutusRJ11
| `--A. c. diversus Bishop 1901RJ11
`--‘Ammodramus’ nelsoni Allen 1875BKB15, RJ11 [=Ammodromus (l. c.) caudacutus nelsoniRJ11]
|--A. n. nelsoniRJ11
|--A. n. alter (Todd 1938)RJ11
`--A. n. subvirgatus Dwight 1887RJ11

*Type species of generic name indicated

References

[B94] Bock, W. J. 1994. History and nomenclature of avian family-group names. Bulletin of the American Museum of Natural History 222: 1–281.

[BF16] Bryson, R. W., Jr, B. C. Faircloth, W. L. E. Tsai, J. E. McCormack & J. Klicka. 2016. Target enrichment of thousands of ultraconserved elements sheds new light on early relationships within New World sparrows (Aves: Passerellidae). Auk 133: 451–458.

[BKB15] Burleigh, J. G., R. T. Kimball & E. L. Braun. 2015. Building the avian tree of life using a large-scale, sparse supermatrix. Molecular Phylogenetics and Evolution 84: 53–63.

Hoyo, J. del, A. Elliott & D. A. Christie (eds) 2011. Handbook of the Birds of the World vol. 16. Tanagers to New World Blackbirds. Lynx Edicions: Barcelona.

Klicka, J., F. K. Barker, K. J. Burns, S. M. Lanyon, I. J. Lovette, J. A. Chaves & R. W. Bryson, Jr. 2014. A comprehensive multilocus assessment of sparrow (Aves: Passerellidae) relationships. Molecular Phylogenetics and Evolution 77: 177–182.

[LG91] Li, W.-H., & D. Graur. 1991. Fundamentals of Molecular Evolution. Sinauer: Sunderland (MA).

[OF19] Oliveros, C. H., D. J. Field, D. T. Ksepka, F. K. Barker, A. Aleixo, M. J. Andersen, P. Alström, B. W. Benz, E. L. Braun, M. J. Braun, G. A. Bravo, R. T. Brumfield, R. T. Chesser, S. Claramunt, J. Cracraft, A. M. Cuervo, E. P. Derryberry, T. C. Glenn, M. G. Harvey, P. A. Hosner, L. Joseph, R. T. Kimball, A. L. Mack, C. M. Miskelly, A. T. Peterson, M. B. Robbins, F. H. Sheldon, L. F. Silveira, B. T. Smith, N. D. White, R. G. Moyle & B. C. Faircloth. 2019. Earth history and the passerine superradiation. Proceedings of the National Academy of Sciences of the USA 116 (16): 7916–7925.

[RJ11] Rising, J. D., A. Jaramillo, J. L. Copete, P. G. Ryan & S. C. Madge. 2011. Family Emberizidae (buntings and New World sparrows). In: Hoyo, J. del, A. Elliott & D. A. Christie (eds) Handbook of the Birds of the World vol. 16. Tanagers to New World Blackbirds pp. 428–683. Lynx Edicions: Barcelona.

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