Pipilo Erythrophthalmus Research Articles

Mitochondrial DNA Phylogeographic Differentiation among Avian Populations and the Evolutionary Significance of Subspecies

ABsTRAcr.-Phylogeographic population structures revealed by restriction analyses of mitochondrial (mt) DNA were assessed within each of six avian species with continentwide distributions in North America. The magnitude and geographic pattern of mtDNA variation differed considerably among species. The Downy Woodpecker (Picoides pubescens) and Mourning Dove (Zenaida macroura) exhibited little mtDNA polymorphism and a shallow phylogeographic structure. The Brown-headed Cowbird (Molothrus ater) and Song Sparrow (Melospiza melodia) showed somewhat higher nucleotide diversity, but no evidence of long-standing population separations. For each of these four species, evolutionary effective sizes of female populations estimated from mtDNA were substantially smaller than population sizes at the present time, suggesting historical demographic constraints on the numbers of females through which mtDNA lineages successfully have been transmitted. In contrast, the Rufous-sided Towhee (Pipilo erythrophthalmus) and Common Yellowthroat (Geothlypis trichas) showed relatively deep mtDNA separations (mean nucleotide sequence divergence p = 0.008 and p = 0.012, respectively) between populations in Washington versus those in the central and eastern states. In the case of the Rufous-sided the mtDNA clades may correspond to morphological and behavioral differences distinguishing the western Spotted Towhee, which was formerly recognized as a distinct species. Overall, however, most of the taxonomic subspecies currently recognized within the six assayed species were genetically very close, and showed no obvious mtDNA differences. These results raise questions concerning the population-genetic and evolutionary significance of current subspecies designations in ornithology. Received 5 June 1991, accepted 23 February 1992.

Seasonal changes in avian song nuclei without seasonal changes in song repertoire

Seasonal variation in the size of song nuclei in the brains of male songbirds may be related to the ability to learn to sing new songs as adults. This hypothesis was tested with the rufous-sided towhee (Pipilo erythrophthalmus), a species in which song repertoires are stable after 1 yr of life. Towhees were hand raised in the laboratory and tutored with normal towhee songs. After song repertoires were recorded at 1 yr of age, photoperiods were manipulated so that 10 male towhees experienced short days and 10 males experienced long days. Circulating hormone concentrations and anatomical attributes of song nuclei were then measured. Photoperiod-related differences in the song nuclei of these towhees were as large as those seen in "open-ended learners" (i.e., species that continue to learn new songs as adults). Seasonal changes of the adult song system may thus occur without disrupting existing song repertoires and without the development of new songs. The synaptic plasticity provided by such seasonal variation, however, may enable song learning by adult birds.

Avian Nesting Ecology in Small Even-Aged Aspen Stands

I examined abundance, location, and success of avian nests from June through August, 1985-87, in small (1 ha) even-aged aspen (Populus spp.) stands on a 240-ha study area managed as ruffed grouse (Bonasa umbellus) habitat in central Pennsylvania. Ninety-five nests of 14 species, mainly rufous-sided towhee (Pipilo erythrophthalmus) (28.4%) and gray catbird (Dumetella carolinensis) (24.2%), were located. Observed versus expected numbers of total nests (all species combined) and those of towhees and catbirds differed (P 0.05) of stand age and distance from an edge but was inversely related (P 0.5 m above ground level) were more susceptible than lower nests (≤0.5 m) to predators

Latent Population Responses of Summer Birds to a Catastrophic, Climatological Event

Catastrophic events can create ecological crunches for avian populations. Ornithologists generally assume that such events elicit immediate, dramatic declines in populations followed by rapid recolonization of a site with habitat recovery. Despite total inundation of habitats within the South Platte floodplain of eastern Colorado during the 1983 breeding season, populations of Brown Thrashers (Toxostoma rufum) and Rufoussided Towhees (Pipilo erythrophthalmus) did not decline significantly from 1982 densities. However, populations of both species declined significantly in 1984 and towhee populations remained low in 1985. These observations support speculations from recent studies of shrubsteppe bird populations that site tenacity may play a stronger role in determining annual densities of breeding birds than previously realized.

Natural occurrence of the nucleopolyhedrosis virus of the gypsy moth,Lymantria dispar [Lep.: Lymantriidae] in wild birds and mammals

Three species of birds and 5 species of mammals were captured in the wild from 2 plots in which mortality from naturally occurring nucleopolyhedrosis virus (NPV) among gypsy moth,Lymantria dispar (L.), larvae was 15% and 70%. Bioassays of intestinal contents showed that blue jays,Cyanocitta cristata (L.), towhees,Pipilo erythrophthalmus (L.), white-footed mice,Peromyscus leucopus (Rafinesque), redback voles,Clethrionomys gapperi (Vigers), raccoons,Procyon lotor (L.), and a chipmunk,Tamias striatus (L.), contained infectious (L.), and masked shrews,Sorex cinereus (Kerr), did not. Comparisons among mice and voles indicated that those collected from the plot in which the NPV mortality was greatest (70%) contained the most virus. We concluded that birds and mammals can passively transport infectious gypsy moth NPV in the wild.

Recognition of conspecific song by the rufous-sided towhee

Dual-speaker experiments conducted with rufous-sided towhees ( Pipilo erythrophthalmus) demonstrated that they do not respond to song of three other species and that the sequence and diversity of phrases within a song, and the time interval between syllables of the note-complex phrase, may be important in species recognition. These results are compared with results of song recognition studies done with species that are extensively sympatric with towhees in eastern North America.

Vocal Dueling among Male Marsh Wrens: Evidence for Ritualized Expressions of Dominance/Subordinance

-Male Marsh Wrens' (Cistothorus palustris) sing with immediate variety (i.e. AB CDE . . .); they progress rapidly through their song repertoires and tend to countersing with same song types. Intensive study of two hand-reared males in laboratory now confirms that both song types and song sequences are learned. Furthermore, leader/follower roles during countersinging duels are not determined at random. In this study, Bird 1 dominated Bird 2 in physical encounters; Bird 2 often followed (i.e. matched) song type just sung by Bird 1, but Bird 1 matched songs of Bird 2 only when songs of latter were electronically amplified. The leader/follower roles in countersinging may be ritualized expressions of dominance and subordinance, respectively, and could reveal to both males and females relative vigor of combatants. Received 20 November 1978, accepted 26 March 1979. IMITATION plays an important role in ontogeny of song in many songbird species (e.g. Nottebohm 1972, Kroodsma 1977), and if juveniles either remain at or return to locality where songs were learned, interacting breeding males will possess similar songs. If a male has only one song type in his repertoire, temporal adjustments in song delivery may be used to achieve interference or avoidance (Wasserman 1977), but when males imitate several songs and develop sizeable repertoires of different song types, potential for complexity of interactions during countersinging is escalated. Studies of Chaffinches (Fringilla coelebs), Great Tits (Parus major), Black-crested Titmice (Parus bicolor), Cardinals (Cardinalis cardinalis), Rufous-sided Towhees (Pipilo erythrophthalmus), etc., have revealed that neighboring males often respond to one another or to a tape recording (of a song from local dialect) with matching song types (Hinde 1958, Gompertz 1961, Lemon 1968a, b, Kroodsma 1971, respectively; see Falls and Krebs 1975 for an apparent exception to this pattern). Males of these species typically sing a song type several times before switching to another (i.e. AAAA . . . BBBB . . .), but potential for vocal interactions is further heightened when successive songs are usually different (i.e. ABCDE . . .), as in Marsh Wren (Verner 1975). With such a rapid interchange of song types, roles of leader and follower in countersinging become immediately evident. Verner (1975: 295) speculated that this song-matching during countersinging reflected the dominance/subordinance relationships between neighbors, but because of complexity of both song repertoire and interactions in field, he could not adequately document relationship among males. Having studied this phenomenon in laboratory with hand-reared birds, I can now (1) confirm that both song types and favored sequences in this complex behavior are learned, (2) reveal that roles of leader and follower in countersinging duels are not determined at random, and (3) support hypothesis that such vocal dueling, together with loudness of delivery, is a reflection of size, age, and/or dominance/subordinance relationships and might be I See Kroodsma and Verner (1978) for encouraged change of common name from Long-billed Marsh Wren to Marsh Wren. 506 The Auk 96: 506-515. July 1979 This content downloaded from 157.55.39.147 on Wed, 18 May 2016 06:05:48 UTC All use subject to http://about.jstor.org/terms July 1979] Vocal Dueling among Marsh Wrens 507

The Relation of Breeding Schedule and Clutch Size to Food Supply in the Rufous-Sided Towhee

Food is important as an ultimate factor in the timing of breeding (Moreau 1950, Lack 1954, 1966, 1968) and in the evolution of clutch size (Lack 1954, 1966, 1968, Cody 1966, 1971, Klomp 1970, von Haartman 1971) in birds. Although early studies emphasized the significance of food for dependent young (see Lack 1954), it now appears that the food available to the laying female may have a proximate effect as well on the date of first egg deposition and on clutch size, at least in some species (Pitelka et al. 1955, Mebs 1964, Perrins 1965, 1970, Cody 1966: Table 1, Lack 1966, Bengtson 1971, Moss et al. 1971). In this paper, I examine the relation of breeding schedule and clutch size to food in populations of Rufous-sided Towhees (Pipilo erythrophthalmus) inhabiting two different habitats along the northeastern Atlantic seaboard.

The Nasal Mites of Birds from Louisiana. VIII. Additional Records and Description of a New Species (Acarina: Dermanyssidae, Ereynetidae, Epidermoptidae, and Cytoditidae)

Sternostoma pirangae sp. n. from the nasal passages of Piranga rubra is described. The new species is most similar to Sternostoma tangarae Fain and Aitken, but differs in having 7 pairs of large cylindrical subapical setae on tarsi II to IV, slightly more rounded podosomal plate which is longer than wide, and 6 solenidions on the sensorial area of tarsus I. New host records are established for Sternostoma lanorium Fain from Lanius ludovicianus, Astrida caprimulgi Fain from Caprimulgus carolinensis, Boydaia spatulata Fain from Sialia sialis, Cytonyssus therae Hyland from Coccyzus americanus, and Turbinoptes strandtmanni Boyd from Tringa solitaria. Also collected were Ptilonyssus Castro from Dendroica pinus, P. japuibensis Castro from Pipilo erythrophthalmus, P. thryothori Pence from Thryothorus ludovicianus, P. hoseini Fain and Aitken from Vireo griseus, P. toxostomae Pence from Toxostoma rufum, Tinaminyssus belopolskii Bregetova from Florida caerulea, Colinoptes cubanensis Fain from Colinus virginianus, Congocoptes furmani Fain from Centurus carolinus, and Cytonyssus troglodyti Pence from Thryothorus ludovicianus. The systematics of the sairae species group of the genus Ptilonyssus is discussed and Ptilonyssus constrictus Ford is regarded as a synonym of P. Castro. Examination of some additional specimens of Louisiana birds has revealed several new and additional host and geographic records for nasal mites and a new species, Sternostoma pirangae. The materials and methods, including the designations used in the following descriptions, have been previously outlined (Pence, 1972a). Family Dermanyssidae Subfamily Rhinonyssinae Genus Tinaminyssus Strandtmann and Wharton Strandtmann and Wharton, 1958, Contrib. Inst. Acar. Univ. Md. 4: 161. Synonymy: Domrow, 1969, Proc. Linn. Soc. N. S. Wales 93: 305. Tinaminyssus belopolskii (Bregetova) Bregetova, 1950, Dokl. Akad. Nauk SSSR 71: 1005. Additional host record: 7 9 9, 1 N from Florida caerulea, little blue heron, Mandeville, La., 15 September 1972, D.B.P. Received for publication 16 April 1973. * Present address: Department of Veterinary and Zoological Medicine, Texas Tech University School of Medicine, Lubbock, Texas 79409. Genus Sternostoma Berlese and Trouessart Berlese and Trouessart, 1889, Bull. Biblioth. Sci. Ouest 2: 128. Sternostoma lanorium Fain Fain, 1956, Rev. Zool. Bot. Afr. 53: 156. New host record: 1 9, 1 8 from Lanius ludovicianus, loggerhead shrike, Mandeville, La., 9 September 1972, D.B.P.

The Nasal Mites of Birds from Louisiana. IV. The Genus Ptilonyssus (Dermanyssidae: Rhinonyssinae) with a Description of Two New Species

A new species and new host records are established for nasal mites of the genus Ptilonyssus from passeriform birds in Louisiana. Ptilonyssus corvi sp. n. is described from Corvus brachyrhynchos. New host records are Ptilonyssus tyrannus from Sayornis phoebe, Contopus virens, and Epidonax virescens, P. callinectoides from Myiarchus crinitus, P. pari from Parus carolinensis, P. hoseini from Vireo solitarius, P. pirangae from Parus bicolor, P. icteridius from Icterus galbula and Euphagus carolinus, P. morofskyi from Geothlypis trichas, Dendroica coronata, D. pinus, Seiurus motacilla, Melospiza georgiana, Pipilo erythrophthalmus, Ammospiza maritima, and Zonotrichia albicollis. In addition, P. vireonis and P. angrensis from Vireo olivaceus and Progne subis, respectively, are reported for the first time from Louisiana. The systematics and host relationships of these species are discussed. Studies on the nasal mites of Louisiana birds have revealed a new species, new host records, and new geographic records for the rhinonyssine genus Ptilonyssus. The present study is concerned with Ptilonyssus sensu lato, or those species which have two dorsal plates, the podosomal and opisthosomal, and chelicerae of uniform diameter or only slightly attenuated at the distal end. The materials and methods, including the designations and abbreviations used in the following descriptions, have been outlined previously (Pence, 1972). Genus Ptilonyssus Berlese and Trouessart Ptilonyssus Berlese and Trouessart, 1889, Bull. Biblioth. Sci. Ouest 2: 128. Synonymy (see Domrow, 1969, Proc. Linn. Soc. N.S.W. 93: 333).

The Nasal Mites of Birds from Louisiana. III. The Genus Ptilonyssus (Dermanyssidae: Rhinonyssinae) with Description of a New Species

A new species and new host records are established for nasal mites of the genus Ptilonyssus from passeriform birds in Louisiana. Ptilonyssus corvi sp. n. is described from Corvus brachyrhynchos. New host records are Ptilonyssus tyrannus from Sayornis phoebe, Contopus virens, and Epidonax virescens, P. callinectoides from Myiarchus crinitus, P. pari from Parus carolinensis, P. hoseini from Vireo solitarius, P. pirangae from Parus bicolor, P. icteridius from Icterus galbula and Euphagus carolinus, P. morofskyi from Geothlypis trichas, Dendroica coronata, D. pinus, Seiurus motacilla, Melospiza georgiana, Pipilo erythrophthalmus, Ammospiza maritima, and Zonotrichia albicollis. In addition, P. vireonis and P. angrensis from Vireo olivaceus and Progne subis, respectively, are reported for the first time from Louisiana. The systematics and host relationships of these species are discussed. Studies on the nasal mites of Louisiana birds have revealed a new species, new host records, and new geographic records for the rhinonyssine genus Ptilonyssus. The present study is concerned with Ptilonyssus sensu lato, or those species which have two dorsal plates, the podosomal and opisthosomal, and chelicerae of uniform diameter or only slightly attenuated at the distal end. The materials and methods, including the designations and abbreviations used in the following descriptions, have been outlined previously (Pence, 1972). Genus Ptilonyssus Berlese and Trouessart Ptilonyssus Berlese and Trouessart, 1889, Bull. Biblioth. Sci. Ouest 2: 128. Synonymy (see Domrow, 1969, Proc. Linn. Soc. N.S.W. 93: 333).

Life Cycle and Development of Prosthorhynchus formosus (Van Cleave, 1918) Travassos, 1926, an Acanthocephalan Parasite of Birds

The life cycle of Prosthorhynchus formosus (Van Cleave, 1918) Travassos, 1926 is presented. The terrestrial isopods Armadillidium vulgare, Porcellio laevis, and P. scaber served as experimental intermediate hosts, and chickens and turkeys as experimental definitive hosts. After ingestion of the eggs by the isopod, the acanthor emerges from its shells within 15 min to 2 hr, enters the gut wall of its host, and remains there 15 to 25 days. It then migrates to the hemocoel and develops through the acanthella stage to the infective cystacanth by the 60th to 65th day of infection. Van Cleave (1918) described Plagiorhynchus formosus on the basis of four specimens collected from the flicker, Colaptes auratus, at Bowie, Maryland. In 1926, Travassos reassigned this species to the genus Prosthorhynchus but gave no reasons for the change. Van Cleave (1942) disagreed strongly with this change in generic status and reaffirmed its position in Plagiorhynchus. Golvan (1956) clarified the differences between Plagiorhynchus and Prosthorhynchus and defended its position in Prosthorhynchus. This view, which was upheld by Petrochenko (1956) and Yamaguti (1963), is accepted by the present authors. Jones (1928) extended the host list to include the chicken (Gallus domesticus), robin (Turdus migratorius), and the crow (Corvus americanus). She remarked on the possibilities of this worm becoming an important parasite of domestic fowl. Cuvillier (1934) added the catbird (Dumatella carolinensis), and a thrush Received for publication 17 April 1964. * Part of a thesis submitted by the senior author to the Graduate School of Colorado State University in partial fulfillment of the requirements for the Ph.D. degree, February 1964. This investigation was supported by Training Grant PHS-2E-94 (C2,3) from the National Institute of Allergy and Infectious Diseases of the National Institutes of Health, U. S. Public Health Service, and by a fellowship from the Boettcher Foundation, Denver, Colorado. (Hylocichla sp.), and Van Cleave (1942) the towhee (Pipilo erythrophthalmus), starling (Sturnus vulgaris), grackle (Quiscalus quiscala), and hermit thrush (Hylocichla guttata). Chandler and Rausch (1949) found it in the brown thrasher (Toxostoma rufum), and Hunter and Quay (1953) in Macgillivray's seaside sparrow (Ammospiza maritima macgillivaraii). The present report adds the red-shafted flicker (Colaptes cafer) and the domestic turkey (experimental). Sinitsin (1929) reported finding a juvenile of Plagiorhynchus formosus in the sowbug, Armadillidium vulgare, near Washington, D. C. To date, this is the only report on the life history of P. formosus, although Dollfus and Dalens (1960) reported larvae of Prosthorhynchus cylindraceus in A. vulgare in France. This work presents the complete life cycle of Prosthorhynchus formosus as observed under laboratory conditions, utilizing three species of terrestrial isopods as intermediate hosts and domestic fowl as definitive hosts. MATERIALS AND METHODS

Some Seasonal Changes in Morphology of the Rufous-Sided Towhee

Seasonal changes have been demonstrated in a number of avian tissues and organs and in bird weight. Such changes indicate not only seasonal changes in morphology but they may, in some instances, suggest the nature of the morphologic and physiological adjustments made by the individual as it passes from one season to the next. This study is concerned with seasonal changes in bill length, in length of the intestinal tract, or gut, and in weight of a resident race of the Rufous-sided Towhee, Pipilo erythrophthalmus megalonyx. Data were derived from samples of males and females collected at, or as close as possible to, two-week intervals near the Hastings Reservation, two and onehalf miles east of Jamesburg, Monterey County, California, from January 5, 1955, to May 10, 1956. Each specimen was autopsied in the field and the gonads were either removed (males) or examined in situ (females). Each bird was weighed on return from the field, its intestinal tract was measured, the amount of stored fat was estimated, and the skin was saved. In addition, the stomach contents were examined and a rough estimate was made of their composition. BILL LENGTH

Ixodes Bishoppi, a New Species from Georgia (Acarina: Ixodidae)

In the course of routine examinations of birds and mammals captured at Savannah, Ga., the authors collected numerous specimens of a new species of tick, Ixodes bishoppi . The species is related to I. minor Neum. and I. muris Bishopp and Smith, and has similar host relationships. The larvae and nymphs were fairly common on cotton rats, Sigmodon hispidus hispidus Say and Ord., and were occasionally found on birds. Of 56 cotton rats examined 40 were infested, carrying a total of 191 larvae and 40 nymphs. Of numerous birds of various species examined 3 were infested; a house wren, Troglodytes aedon aedon (Vieill.), with 5 larvae and 1 nymph, a house wren with 8 larvae, and a towhee, Pipilo erythrophthalmus erythrophthalmus (Linn.), with 6 larvae and 2 nymphs. One of several house mice, Mus musculus musculus Linn., caught in a grassy field was infested with 2 larvae. Of 13 cottontail rabbits, Sylvilagus floridanus ssp., only 1 was infested, carrying a single nymph. Adult ticks were collected only twice, in each instance from a cotton rat. One rat was infested with 1 male and 7 females, the other with a single female. Numerous males and females, including the types, were reared from specimens collected as larvae or nymphs.