This dissertation has been microfilmed exactly as received bu-loUo

MAXWELL n, George Ralph, 1935- LIFE HISTORY OF THE COMMON GRACKLE, Quiscalus quiscula (Linnaeus).

The Ohio State University, Ph.D., 1965 Zoology

University Microfilms, Inc., Ann Arbor, Michigan LIFE HISTORY OF THE COMMON GRACKLE, Qulsoalus gulsoula (Linnaeus)

DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Dootor of Philosophy in the Graduate School of The Ohio State University

By

George Ralph Maxwell II, B.A., M.S.

The Ohio State University 19 65

Approved "by

Li Adviser Department of Zoology and Entomology Dedicated to Susan, for her help, understanding, and for the sacrifices which she has made during the years of my graduate training.

ii Acknowledgments

The author wishes to express sincere appreciation to his dissertation adviser, Professor Loren S. Putnam, for his friendly counsel, direction and efforts to achieve a good working situation necessary for productive re­ search. I am also indebted to the Department of Zoology and Entomology of The Ohio State University for providing logistical support, especially during the data gathering phase at The Franz T. Stone Laboratory, Put-in-Bay, Ohio. Thanks are also extended to the following, who helped in so many ways to bring this projeot to completion* Dr. W.H. Anderson, Agricultural Research Service, Belts- ville, Maryland; Dr. W.T. Atyeo, University of Nebraska, Lincoln, Nebraska; Mr. Earl B. Baysinger, Migratory Bird Populations Station, Laurel, Maryland; Mr. John Condit, Eastmoor High School, Columbus, Ohio; Dr. John L. Crites, The Ohio State University, Columbus, Ohio; Mr. Thomas Duff, Put-in-Bay, Ohio; Dr. M.L. Giltz, The Ohio State University, Columbus, Ohio; Dr. E.E. Good, The Ohio State University, Columbus, Ohio; Mr, Donald E. Johnston, Institute of Aearology, Wooster, Ohio; Dr. Mildred Mlsklmen, Rutgers university, New Brunswick, New Jersey; Mr. Steve Putnam, Worthington, Ohio; Mr. Thomas C. Rambo, The Ohio State University, Columbus, Ohio; Mr. Everett Seymour, Put-in-Bay, ill Ohio; Dr. Russell Skavaril, The Ohio State University, Columbus, Ohio; Mr. Hussell Smith, Put-in-Bay, Ohio; Mrs. Elizabeth Snider, The Ohio State University, Columbus, Ohio; Dr. R.W. Strandtmann, Texas Technological College, Lubbock, Texas; Dr. and Mrs. Milton Trautman, The Ohio State university, Columbus, Ohio; Dr. G.W. Wharton, The Ohio State University, Columbus, Ohio. I am grateful to the National Science Foundation for their financial assistance througn a summer Fellow­ ship in 1964 and to The Ohio State University for a Muellhaupt Fellowship in 1964 and an Osbum Fellowship in the summer of 1965. Without this assistance my work could not have been completed in three years.

iv Vita

27 March, 1935 B o m - Morgantown, West Virginia 1957 . • . B.A., West Virginia University, Morgantown 1958-1960 . Teaching Assistant, Dept, of Biology, West Virginia University, Morgantown 1961 . , , M.S., West Virginia University, Morgantown 1961-1962 . Instructor in Preventive Medicine, Medical Field Service School, Fort Sam Houston, Texas 1962-1964 . Teaching Assistant, Dept, of Zoology, The Ohio State University, Columbus 1964 . . . N.S.F. Fellow, Franz T. Stone Laboratory, Put-in-Bay, Ohio 1964-1965 . Muellhaupt Fellow, Dept, of Zoology, The Ohio State University, Columbus

1965 • • • Osbura Fellow, Dept, of Zoology, The Ohio State University, Columbus

Publications

Neutralization of ten snake venoms by homologous and heterologous antivenins. H.L. Keegan, F.W. Whitt©more and G.R. Maxwell. Copeia, No. 2, pp. 313-31 6 , July, 1962. Wing pad and tergite growth of mayfly njtmphs in winter. G.R. Maxwell and A. Benson. Amer. Mid. Nat., Vol. 6 9 , pp. 224-230, January, 1963*

Fields of Study

Major Field: Vertebrate Zoology and Ornithology Contents

Page Introduction 1

The Study Area 3 Materials and Methods 5 Taxonomy 7 Basis for identification 7 History of generic and specific classification 10 Formal description of Ouisoalus auisoula (Linnaeus) 14 Synonomy 1**

Diagnosis 17 Description 20 Geographic variation 21

Individual variation 25 Sexual variation 30

Ontogenetic variation 31 Distribution 33 Recognition of related forms 36 Voice 41 Courtship Behavior 46 Pair formation 46 Sexual behavior 50 Pag© The Nest 51 Nest anatomy 51 Looation of the nest 52 Nest -building 53 Territory 57 Eggs 60 The egg 60 Egg laying 66 Clutoh size 69 Inoubation 71 Length of the inoubation period 87 Care of the Young 91 Brooding 91 Feeding 98 Nest maintenance 10*1-

Fledging and Post-nesting Activities 110 Nest Success 113 Development of Young 117 Migration 153 Spring and Autumn 153 Winter range 155 Longevity 161

vll Page Pood Habits 162 Introduction 162 Adult food types 162 Nestling food types 171 Unusual feeding behavior 173 Parasitism 177 Summary 189 Appendix 197 Bibliography 212

viii Tables

Table Page 1. Comparative feather ooloratlon 22 2. Body measurements of specimens of £•£• qulsoula 23 3. Body measurements of specimens of stonel 23 4. Body measurements of specimens of £•

13. Egg laying times 67 14. Clutch size 70 15. Inoubation patterns 77 16. Attentive behavior during the normal Incubation period 80 17. Attentive behavior during the abnormal Inoubation period 81 18. Summary of data for Inoubation patterns 85 19. Length of the inoubation period 88 ix Table Page

20. Number of feeding visits in relation to time of day 105

21. Nesting sucoess 115 22. Causes of nest failure 115

23. Daily weights of nestlings 138 24. Dally length of nestling body parts 142

25. Growth rates in mm/day of nestling body parts 143 26. Counts of Common Grackles at extremes of winter range 156

27. Areas with highest ooncentration of Common Grackles in winter 158 28. Areas of recoveries of Ohio banded graokles 160

29. Weights and types of stomach contents 165 30. Pood of the graokle by month 168

31. Identified insects from stomachs of the graokle 169 32. Food types of nestling Common Graokles 174

33. Daily morning and afternoon temperatures 198

3^« Measurements of Q.a. versioolor eggs 1Q9 35. Summary of feedings by adult grackles at Nest 8-64 206 36. Disposition of time by the female at two nests 207

37. Brooding and feeding of nestlings at Nest 8-64 208 38. Brooding and feeding of nestlings at Nest 20-64 209

39. Duration of care of nestlings at Nest 8-64 210 40. Duration of care of nestlings at Nest 20-64 211 Figures

Figure Page 1. Map of distribution of the Common Graokle 34 2. Inoubation patterns at Nest 20-64 74 3. Inoubation patterns at Nest 6-64 75 4. Effect of air temperature on constancy 83 5. Brooding sessions per hour at Nest 8-64 93 6 . Disposition of females' time at Nest 8-64 95 7* Disposition of females' time at Nest 20-64 96 8 . Comparison of brooding visits at Nests 8-64, 20-64 97 9 . Comparison of feeding visits at Nests 8-64, 20-64 100 10. Comparison of the number of feeding visits by the male and female at Nest 8-64 101 11. Comparison of the number of feeding visits by the male and female at Nest 20-64 102 12. Nest sanitation by the male and female at Nest 8-64 107 13. Nest sanitation by the male and female at Nest 20-64 108 14. One day old nestling 118 15• Two day old nestling 119 16. Three day old nestling 120 17* Four day old nestling 121 18. Five day old nestling 122 19. Six day old nestling 123 20. Seven day old nestling 124

xi Figure Page 21. Eight day old nestling 12 5 22. Nine day old nestling 126 2 3 . Ten day old nestling 12? 24. Eleven day old nestling 128 2 5 . Twelve day old nestling 129 26. Age-weight relationships of nestlings from Nest 8-64 137 2 7 . Average growth of nestlings from Nest 8-64 140 28. Ventral view of major feather tracts 145 2 9 . Dorsal view of major feather tracts 147 30. Lateral view of major feather tracts 149

xll Introduction

Many members of the family Icteridae have been the subject of recent life history and behavioral studies due, in part, to their rapid population increases over most of North America. As the population expands, their economic importance becomes more significant, which lnoreases the need for basic information about their life prooesses. The Common Graokle1 (Qulsoalus quisoula versicolor) is represented in recent literature by one published life his­ tory paper (Bent, 1958) and one unpublished paper (Eyer, 195*0 • Two other reports of significance are a nesting study by Petersen and Young (1950) and a behavior study by Ficken (1963). Most of the other information is scat­ tered and in brief form in Journals. This dissertation was undertaken with the aim to present new knowledge in the areas not discussed in the limited publications available and to add additional data on subjects previously presented. This paper contains three major areas of investigation* a taxonomio review, the nesting cycle and a parasitological section.

*-The common name for this subspecific form, Bronzed Grackle, is not used throughout this paper in an effort to minimize the effect of nomenclatorial "splitting" during past years. The taxonomic section covers the history of the nomen­ clature, a formal description, variation, distribution, and recognition of related forms. The nesting cyole consists of a discussion of voioe, pair formation, territory, the nest, the eggs, incubation, care of the young, development of the young, post-nesting activities and migration. The parasite section Is not intended to be a complete list of grackle ecto-and endoparasites, but rather a summary of those species found in or on graokles I processed, supplemented by the results of a brief liter­ ature survey. The material has been organized in chronological order, when such order is necessary for an understanding of the sequence of events. The data were collected and assembled during the period from March, 1964 to July, 19^5• The major portion of the field data was collected at The Franz T. Stone Laboratory, Put-in-Bay, South Bass Island (Lake Erie), Ottawa County, Ohio, and literature supplement was obtained on The Ohio State University campus in Columbus, Ohio. Th* Study Area

During the spring and summer of 1964 and the spring of 1965, this study was completed on South Bass Island, one of the three major islands in the Bass Islands Archipelago in western Lake Erie, Ottawa County, Ohio. Put-in-Bay, a summer resort, is located on this Island, which Is approximately five miles north of the Ohio mainland and five miles south of the Canadian border. The study area was specifically In the Peach Point peninsula, a small extension of land from the north-central part of the island. Peach Point is a park type area consist­ ing of lawns, gardens and orchards. The red cedar (Junlperus vlrglnlana) is the dominant tree on the lawns of the penin­ sula and the tree used by the Common Graokle for nesting. Beecher (1942) studied avlan-plant communities in Lake County, Illinois, and found the graokle to breed in willow rows in low prairie. Eyer (1954) found the graokles to build nests in Spiraea bushes in a marsh near Lansing, Michigan. I found grackles building in buttonbush (Cephalanthus oooidentalls) in a marsh on North Bass Island, Ottawa County, Ohio. The following birds were found to nest in association with the graokle on South Bass Island: Bobin (Turdus migra- torlus). Mourning Dove (Zenaldura macroura), Kingbird (Tyrannus tyrannus). Yellow Warbler (Dendroloa aestlve),

House Wren (Troglodytes aedon), Indigo Bunting (Passerlna oyanea), Goldfinch (SPlnus trlstls). Song Sparrow (Melospiza melodla), Redstart (Setojghaga rutlcllln), Red-eyed Vlreo (Vlreo ollvaoeus), Red-wing (Agelalus phoenlceus)« and Cedar Waxwing (Bombyollla oedrorum). Materials and Methods

Data collection for this paper has been largely a matter of field observation. As many phases of the life history were studied as possible, but the emphasis is upon the characteristics of the breeding population. The equip­ ment necessary for this type of study is simple, consisting of 7-power binoculars, 15-power spotting telescope, blinds, ladder, pole-mlrror and a note book. The blinds were per­ manent structures, either houses, a oar, or natural vege­ tation. The ladder and pole-mirror were necessary for observations at nest level. The pole-mirror consisted of a by 6 inch mirror attached at a right angle to a 10 foot pole. This was a very useful tool for determining egg lay­ ing and hatching times. The development of the nestlings study required a knife edge balance, 35 bub* camera, calipers, millimeter rule and color bands. The young birds were weighed to the nearest l/10th of a gram, twice a day and four body parts measured once a day. Each morning the nestlings were photographed on a piece of white paper for the duration of the nestling period.

Parasites were oollected from freshly killed graokles by washing them in 70 percent alcohol and pipetting the settled organisms into a watoh glass for binocular mioroscope observation. The parasites were then mounted, on slides and labeled. Identification was achieved by sending the slides to applicable specialists. The nomen­ clature was checked by the Institute of Aoarology at Wooster, Ohio. The intestine was out open throughout its entire length and examined under the binocular microscope. Attempts were made to color band adults so that individual recognition could be made. Baited traps with a small opening for an entrance were used with limited success. I was not able to band both members of one pair at nests under study, but managed to get bands on six adults associated with area nests. The bands were colored plastic and sealed on the bird*s leg with xylene. Two broods of nestlings were color banded before they fledged. The food study required fresh stomachs obtained from graokles killed with a smooth-bore .22 caliber collection rifle. The stomachs were cut open, emptied into a watch glass and washed with water. The materials were sorted

into plant and animal portions, then dried for Zb hours in a oonstant temperature oven. Dried remains were weighed on an analytical balance, sorted for identifi­ cation purposes and recorded. Taxonomy

Basis for Identification The grackles studied in the field and museum were Identified by comparison with keys or published descrip­ tions of the species and/or subspecies. These methods of Identification proved them to be Qulscalus qulsoula (Linnaeus)• The characteristics for each taxon are placed in paragraphs followed by oitations to the source. The following characteristics place the grackles in the order Passeriformes. Toes four, three in front, one behind, cleft to the base or there immovably coherent; hind toe not elevated; nostrils not opening beneath soft swollen membrane; bill not cered and hooked; second­ aries more than six; primaries nine or ten (Coues, 1903). These characteristics placed the grackles in the family Ioterldae. Toes, three in front, one behind? hind toe nbt inserted above the level of the rest (and generally but not always not shorter than the shortest front toe); primaries nine, the first of variable length; bill oonirostral, usually thick, stout, and with evident angulation of the commissure (Coues, 1903). Tarsus longer

7 than middle toe with claw and compressed to a sharp ridge behind, with a longitudinal groove on Its outer surface; tall at least 2/3 length of wing; primaries nine, outermost more than 1/2 length of longest, longest primary less than twice length of Inner primary or secondaries; length of mandibular ramus more than 1/2 length of gonys; bill stout, conlrostral and unhooked; nostrils oval or slitlike and Imperforate; without riotal bristles (Blair et al., 1957). The following are key characters of the genus Qulscalus. Three to five primaries with outer webs sinuated, ninth primary not shorter than third, usually longer, longest tertials not produced beyond secondaries; reotrioes not acuminate, tail plicate (V-shaped) and graduated; mesorhinum very broad, one-fourth as wide as length of oulmen; nostrils with more or less dlstlnot superior operculum or membrane, large, narrow, more or less linear, oblong, or subouneate well forward of the loral antlae; oulmen more or less strongly and abruptly deourved terminally or else culmen less than 1/3 as long as tall, the latter longer than wing and much graduated; median palatal ridge truncated, angulated, and highest anteriorly; middle toe with claw little if any longer than tarsus, usually shorter; middle toe with claw shorter than its terminal phalanx; hallux with its claw shorter than the digit, not longer than lateral anterior toes; middle phalanx of middle toe shorter than terminal phalanx; outer toe (without claw) reaching to or beyond second (subterminal) Joint of middle toe (Ridgway, 1902). More than two primaries sinuate, out primary shorter than second; feet not reaohing end of tall; oulmen straight or curved; tail graduated for 1/3 its length or more (Blair et al., 1957). Characteristics of the species Qulsoalus qulsoula (Linnaeus), Common Grackle are as follows: blackish; head and neck glossed with purple or greenish blue; wings and tail glossed with purple; body glossed with plain bronze or squamate with bronze green and purplish; iris pale yellow; bill and feet black (Blair et al., 1957). The subspecies of the Common Grackle have, according to Ridgway (1902), these characteristics: Purple Graokle: larger, except bill and feet; wing of adult male averaging 1^3.8 mm., tall 135.6 mm.; adult female wing 127.8 mm., tail 112 mm.; Individual variation in plumage very great. Florida Graokle: smaller, except bill and feet; adult male averaging in the wing 133*1 mm., tall 121.1 mm.; adult female wing 119 mm., tail 106.2 mm.; individual variation in plumage slight. Bronzed Grackle: larger, plumage of back, scapulars, rump, and under parts of body perfectly uniform bronze or brassy olive; lesser and middle wing-coverts entirely bronzy purple or purplish bronze. 10 History of generic and speolflo classification The nomenclature of the Common Graokle represents one of the most confused, but Interesting nomenolatorlal problems In avian taxonomy. This historical account will discuss name changes and the present status of Qulsoalus Qulsoula (Linnaeus). Hellmayr and Conover (1937) cite five generic names for the graokle: Qulsoalus. Qulscala. Chalcophanes. Soaphldurua and Soaphura. The latter four are synonyms of Qulsoalus. Linnaeus (1758) first described the species and named it Graoula qulsoula. This Is the type of the Florida Grackle, Qulsoalus qulsoula qulsoula (Linnaeus). The new generic name Qulsoalus was proposed by Vieillot in 1816 by subsequent designation (Solater, 1886). Sclater (1884) cites three subspecies of Qulsoalus versloolor: Qulsoalus versicolor typious. £.V. aeneus. S*Z* aglaeus. Chapman (1892) uses the specific name qulsoula In place of versloolor. The three subspecies have variously ohanged. The status of the Purple Graokle and Florida Graokle are most confused due to a misldent- ifioatlon of the original type Graoula qulsoula Linnaeus. The nomenolatorlal changes of the Bronzed Grackle are not so complicated. 11

The second edition of the A.O.U. Cheok-list (1895) cites Qulsoalus qulsoula (Linnaeus), Purple Graokle as a separate species and the following two subspecies; Qulsoalus qulsoula aglaeus (Baird), Florida Graokle, and Qulsoalus qulsoula aeneus (Rldgwa$, Bronzed Graokle. Ridgway (1902) used the subspeolflo name qulsoula to denote the Purple Graokle as a subspecies, Qulsoalus qulsoula qulsoula. Wayne (1918) discovered that the type of the Purple Grackle, Graoula qulsoula of Linnaeus, based ohiefly on Catesby's (1731) Monedula purpurea, was In fact a Florida Graokle, Qulsoula qulsoula aglaeus. This relegates Qulsoalus qulsoula aglaeus (Baird) to synonymy, since the latter name applies to the same bird as Graoula qulsoula Linnaeus. All previous names for the Purple Grackle thus become synonyms for the Florida Graokle, Qulsoalus qulsoula qulsoula (Linnaeus). These changes leave the Purple Graokle without a name. Oberholser (1919) in agreement with Wayne's (1918) views recognized the need of a new name for the Purple Graokle. Using a bird from Washington, D.C., as the type, he proposed Qulsoalus qulsoula rldgwavi for the Purple Graokle. Chapman (1935) found that the type from Washington, D.C., was not a true Purple Graokle, but a hybrid between the latter and the Bronzed Graokle. He felt the ridgway1 subspecies should be retained for the hybrid, but the fifth edition of the A.O.U. Check­ list (1957) does not cite this subspecies. It Is doubt­ ful that a hybrid between two subspecies can attain subspeolflo status. Chapman (1935) chose a specimen of the true Purple Graokle from Lakehunbt, N.J., and designated It as the type of Qulsoalus qulsoula stonel Chapman, Purple Graokle. This name Is cited In the fifth edition of the A.O.U. Check-list (1957). The nomenclature of the Bronzed Graokle had remained relatively stable until 1939. Its scientific name Qulsoalus oulscula aeneus was based on Bidgway (1902)• The only major change previous to this time was the use of the name versloolor by Sohlater (1884-). This was changed to qulsoula by Chapman (1892). tfetmore

(1939m) examined Vielllot*s type of Qulsoalus versloolor In the Paris National Museum of Natural History and found it to be the Bronzed Grackle and not the Florida Graokle as previously thought. Chapman (1959) argues against this change on the basis that the Paris Museum speoimen was Improperly labeled, and Vieillot's descrip­ tion fits the Florida Graokle. Wetmore (1939^) defends his observations made at the Paris Museum and states that It Is necessary to accept the labeled specimen as the type of Qulsoalus versloolor. Wetmore (1939a) gives the Bronzed Grackle a species status, Qulsoalus versioolor. In the Twenty-third Supplement to the Fourth Edition of the A.O.U. Check-list (19^8) the Bronzed Grackle has again been given a subspeoifio status, Qulsoalus qulsoula versioolor. In summary, the Fifth Edition of the A.O.U* Check­ list (1957) oites one speoiest Qulsoalus qulsoula (Linnaeus), Common Grackle, and three subspeciest Qulsoalus qulsoula stonel Chapman, Purple Grackle; Qulsoalus qulsoula qulsoula (Linnaeus), Florida Grackle; and Qulsoalus qulsoula versioolor Vieillot, Bronzed Graokle. 14 Formal Description of Qulsoalus qulsoula (Linnaeus)

Synonomy

Genus Qulsoalus Vielllot Qulsoalus Vlelllot. 1816. Anal. Nouv. Om. Elem., p. 3 6 , Type, "by subs, desig. (Gray, List Gen. Bds., p. 41, 1840), Graoula "qulsoala*1 (»»qulsoula) Linnaeus. Qulsoala Lichtenstein, 1823. Verz. Doubl. Berliner Mus., p . 18. Type, by monotypy, Gracula qulsoula Linnaeus. Chaloophanes Wagler, 1827. Syst. Av., I, Fol. 20, Gen. Graoula. spec. 3,4. Type, by subs desig. (Solater, Ibis, 1884, p. 153)» Qulsoalus versicolor Vlelllot* Graoula qulsoula Linnaeus. Soaphldurus Swalnson, 1837. Phil. Mag., (n.s.), I, No. 6 , p. 437. New name for Qulsoalus Vielllot. Soaphura Gloger, 1841. Gemeinn. Hand-und Hllfbuch Naturg., I, p. 26l. New name for Qulsoala Lichtenstein.

Qulsoalus qulsoula qulsoula (Linnaeus) Graoula qulsoula Linnaeus. 1758. Syst. Nat., 10th ed,, I, p. 109. Based on Monedula purpurea Gatesby, Nat. Hist. Carolina, 1731> I» P* 12, pi. 12; South Carolina.

Orlolus ludovlolanus Gmelln. 1788. Syst. Nat., I, 1, p. 387. Based on "Cassique de la Louislane", Buffon, Hist. Nat. des 01s., Ill, p. 242; Planch. Enlum., No. 646, (partial albino); Louisiana. Stumus qulsoala Daudin, 1800. Traite Elem. et Compl. d'Oraith., II, p. 316. New name for Gracula qulsoula Linnaeus. Graoula qulsoala Wilson, 1811. Amer. 0mlth., Ill, p. 44, pi. XXI, fig. 4 (=Graoula qulsoula Linnaeus). Qulsoala nitens Lichtenstein, I823. Verz. Doubl. Berliner Mus., p. 18. New name for Graoula qulsoula Linnaeus. 15 Qulsoalus purpureus Stephens, 1826. Shaw Gen. Zool., XIV, (1), p. 48. New name for Graoula qulsoula Shaw (Gen. Zool., Vii, (2), p. 458, 1809) and Graoula qulsoala Wilson (Amer. Orn., Ill, p. 44, pi. Si, fig. 4, 1811) = Graoula qulsoula Linnaeus. §ulsoalus purpuratur Swalnson, 1837. Anlm. Menag., ; North America (Type now In University Museum, Cambridge, Engl.; of. Sclater, Ibis, 1884, p. 154). Qulsoalus agaeus Baird, 1866. Amer. Journ. Sci., (2), XLI, p. 84. Based on Qulsoalus barltus (not Graoula barlta Linnaeus) Baird, Rep. Expl. Surv. Pacif. R.R., IX, p. 556, 1858; Key Bisoayne, Cape Florida, Florida (type in U.S. National Museum); Ridgway, Proo. Aoad. Nat. Soi. Phila., 1869, P. 135» southern Florida. Qulsoalus versioolor aglaeus Sclater, 1884. Ibis, p. 154; Florida. Qulsoalus qulsoula Chapman, 1892. Bull. Amer. Mus. N.H., IV, p. 3. Qulsoalus qulsoula aglaeus Chapman, 1892. Bull. Amer. Mus. N.H., IV, p. 5* Qulsoalus qulsoula qulsoula Ridgway, 1902, Bull, U.S. Nat. Mus., L, part 2, p. 215. Qulsoalus versioolor typlous Ridgway, 1902. Bull. U.S. Nat. Mus., L, part 2, p. 217. Qulsoalus qulsoula qulscula Wayne, 1918. Auk, XXXV, p. 440.

Qulsoalus qulsoula qulsoula Oberholser, 1919, Auk, XXXVI, p. 549. 16

Qulsoalus qulsoula stonel Chapman

Qulsoalus qulsoula stonel Chapman, 1935. Auk, LII, p. 25; Lakehurst, N.J. Type In Amer. Mus. Nat. Hist., no. 99687, June 8, 1907.

Qulsoalus qulsoula versioolor Vielllot

Qulsoalus versioolor Vielllot, 1819. Nouv. Diet. Hist. Nat., nouv. ed., XXVIII, p. 488; Etats-Unis. Qulsoalus aeneus Hldgway, I869. Proc. Acad. Nat. Sci. Phlla., XXI, (2), p. 13^. Type, from Mount Carmel, Illinois, in U.S. National Museum; Chapman, Bull, Amer. Mus. Nat. Hist., IV, p. 3» 1892. Qulsoalus versioolor aeneus Solater, 188^. Ibis, p. 15*n Qulsoalus versioolor aenea Solater, 1886. Cat. Bds. Brit. Mus., XI, p. 395. Qulsoalus qulsoula aeneus Ridgway, 1902. Bull. U.S. Nat. Mus., L, part 2, p. 219; Oberholser, Auk, XXXVI, p. 55^» 1919. Qulsoalus versioolor Wetmore, 1939. Proo. U.S. Nat. Mus., LXXXVI, p. 230. 17

Diagnosis Qulsoalus qulsoula is a medium-sized or rather large semi-terrestrial Ioteridae, with long, graduated, and plioate tall; anteriorly truncated and very prominent median palatal ridge; bill about as long as head and strongly decurved at tip; oolor black, with various and strongly contrasted metallio (green, blue, violet and bronze) hues. Bill about as long as head or a little longer, elongate-conical, but upper outline decidedly convex terminally and decurved at tip, its basal depth less than one-half the length of oulmen, but equal to or more than one-half the distance from nostril to tip of maxilla, the basal width a little less; culmen nearly straight (sometimes faintly depressed) in middle, convex terminally, with strongly deourved tip, elevated and usually slightly arched basally, distinctly ridged but the ridge broad and rounded; gonys straight or nearly so, sometimes faintly concave terminally, sometimes faintly convex basally, slightly shorter than maxilla from nostril; maxillary tomium slightly oonvex in middle, slightly concave anteriorly and subbasally, strongly deflexed from beneath nostril to rictus; mandibular tomium oonvex terminally, straight or slightly concave in middle, more or less oonvex and elevated opposite pal­ atal ridge, then slightly deflexed for a short distance, strongly deflexed from beneath nostril to rictus. Nostril more or less triangular (obtusely pointed anteriorly), posteriorly in contact with feathering of loral antiae, overhung by a rather broad membraneous or suboorneous operculum. Wing moderate (less than four to more than four times as long as oulmen, three and a half to nearly four times as long as tarsus), the tip moderately pro­ duced (about equal in length to tarsus or Intermediate between length of tarsus and length of oulmen), sub- truncate; ninth (outermost) primary intermediate between seventh and sixth, between sixth and fifth, or equal to fifth; eighth and seventh or eighth, seventh, and sixth primaries longest; inner web of three outer primaries slightly but distinctly sinuated, but the middle portion not conspicuously widened. Tall plicate, shorter than wing (the difference equal to about one-fourth to one- third the length of culmen), graduated for a little less than length of oulmen, the reotrioes (except middle pair) widest terminally and with obliquely subtruncated tips, the inner web usually a little longer than the outer. Tarsus slightly longer than oulmen, with anterior scutella distinct; middle toe, with claw, decidedly shorter than tarsus; lateral toes with claws falling decidedly short of base of middle claw; hallux equal in length to lateral toes, much shorter, its claw decidedly shorter than the digit; claws only moderately curved (Ridgway, 1902). 20

Description Mult male. Head, neck, and chest varying in color from metallic violet to bronze green; prevailing color of back and scapulars varying from bronzy purple or polished bronze to metallic olive-green or bottle green; rump varying in color from purplish bronze to violet, the color usually more or less broken by admixture of other metallic hues; prevailing color of wings varying from violet-purple to steel blue; tail dark purple, violet, blue, or green, or, in worn or faded plumage, black, glossed with one of these colors; under parts metallic purple, violet, blue, green, etc., the color varying in different parts, sometimes mixes with golden bronze; bill, legs, and feet black; iris pale yellow or yellowish white. Mult female. Decidedly smaller than male and much duller in color, the metallic hues more subdued, sometimes very faint. Young. Uniform sooty, rather paler below, where sometimes showing indistinct streaks of darker (Ridgway, 1902). Range. Eastern North America, west to base of Rockies, in Montana, Wyoming, Colorado, and New Mexico. Winters from Delaware and Ohio Valleys southward (Blair

et al., 1957). 21

Geographic variation The geographic variation in the speoles is restricted to size and feather color differences. Chapman (1935) reoognlzes size and oolor differences which form the basis for the three subspecies. The feather colors characterizing Qulsoalus qulsoula qulsoula - the Florida Grackle, and Quls­ oalus qulsoula versicolor - the Bronzed Graokle, are stable within the subspecies except where they intergrade with Qulsoalus qulsoula stonel - the Purple Graokle. The latter subspecies* oolor differences are so unstable that three phases are oited by Chapman (1935)• See Table 1 for compar­ ison of body parts coloration in the three subspecies. Some size variation among the subspecies is noted from measurements of specimens in the Ohio State Museum and from the literature. The Florida Grackle is the smallest subspecies with the wing and tall lengths muoh shorter than the Purple or Bronzed Graokles. The latter two subspeoles are very similar in size (Tables 2,3 and TABLE 1

Comparative feather coloration in the species Quiscalus quiscula (Chapman, I892).

Feather Subspecies Location Florida Grackle Bronzed Grackle Purple Grackle Q.q. quiscula Q.q. versicolor phase 1 phase 2 phase 3 Q.q. stonei

Head, neck shining dark shining greenish to steel blue purple, steel same as phase 2. and chest violet, bluish purple blue, rarely blue or anteriorly and bluish violet. steel green. bronzy posteriorly.

Back bottle-green, the uniform brassy same as widely margined margined with feathers with a bronze to olive- Q.q. quiscula with bronzy brassy bluish- concealed bronze, no purple, an green, iridescent iridescent bar at iridescent or iridescent band band less clearly the base. other markings. with concealed defined. base brassy bronze to bronzy purple.

Rump purplish or violet like back. same as bronze, brassy brassy bronze with washed with bronzy Q.q.quiscula bronze or bronzy purplish reflections and with iridescent purple. some with terminal spots near ends of iridescent spots. some feathers.

Tail greenish or bluish purplish black same as same as phase 1 same as phase 2. black Q.q. quiscula but glossed.

Wing primaries blackish primaries black,inner same as same as phase 1 same as phase 2. green, inner quills quills violet, bronze Q.q. quiscula but glossed. to purplish violet, tinged, no iridescent lesser coverts with markings. iridescent tips. 23

TABLE 2

Measurements in mm. of ten male and ten female specimens of Quiscalus quiscula quiscula from Ridgway (1902). Note size differential between males and females in percent.

Body Part Range (d) Range (9) Range (d9) Mean (cf) Mean (9) d 9 size differential (d larger)

Wing 129.0-135.9 116.6-121*. 5 116.6-135.9 133.1 119.1 11$

Tail 116.1-128.5 100.8-111.0 100.8-128.5 121.2 106.2 13%

Tarsus 36.6-37.3 33.3—3U.8 33.3-37.3 36.6 33.8 8%

Culmen 31.0-35.1 29.2-31.2 29.2-35.1 33.3 30.2 9%

TABLE 3

Measurements in mm. of ten male and five female specimens of Quiscalus quiscula stonei from Ridgway (1902). Note size differential between males and females in percent.

Body Part Range (d) Range (9) Range (d*9) Mean (d) Mean (9) d $ size differential (cf larger)

Wing 137.7-1U6.8 122.U-133.9 122. U—11*6.8 11*3.8 127.8 12$

Tail 129.5-139.7 101.6-122.7 101.6-139.7 135.6 112.0 18$

Tarsus 31*.3-38.1* 32.8-36.1 32.8-38.1* 36.6 31*.3 7$

Culmen 32.5-36.3 27.9-30.2 27.9-36.3 33.8 29.2 H*$ 24-

TABLE 4

Measurements in mm. of thirty-two male and nine female specimens of Quiscalus quiscula versicolor from the Ohio State Museum. Note size differential between males and females in percent.

Body Part Range (cf) Range (9) Range (d9) Mean (d) Mean (9) d 9 size differential (d larger)

Wing 131.0-148.1 125.0-130.0 125.0-148.1 142.0 127.5 11%

Tail 93.5-138.$ 100.0-116.0 93.5-138.5 124.0 108.7 13%

Tarsus 31.0-37.0 30.5-32.0 30.5-37.0 34.3 31.6 8%

Culmen 28.0-36.0 28.0-32.0 28.0-36.0 32.9 30.2 9% 25 Individual variation Measurements of Individual wing and tall feathers were made on nine male and seven female adult specimens taken at Put-in-Bay, Ohio, in July, 1964. The feathers were measured from the superior umbilicus to the tip of the rhaohis. Tables 5 and 6 give a summary of these measure­ ments. An analysis of variance showed that individuals varied significantly with respect to wing and tail feather lengths. Also the lengths of the individual wing and tall feathers on one specimen differ significantly from the others. Tables 7 and 8 give the statistical analysis of these measurements. The sixth and seventh primaries (primary feather no. 1 is proximal, primary feather no. 9 is distal) are the longest wing feathers, the sixth in the male being slightly longer than the seventh. The first secondary is the longest, the remaining secondaries decrease in size to the smallest or no. 9 secondary. The central tail feathers (feather no. 1) are longest in both sexes, with the gradually decreasing lateral feathers forming a distinct wedge shape or plicate tail. Harmon (1939) reported collecting a Bronzed Graokle near Oklahoma City with the five middle tall feathers white.

I collected a graokle at Put-ln-Bay, Ohio, with one white breast feather. 26

TABLE 5

in mm. of male Quiscalus quiscula versicolor wing -hers from Put-in-Bay, Ohio.

Individual Specimens ABCDEFGHI Average

9 101 101* 106 98 101* 107 102 103 101 102.9

8 103 107 108 103 106 107 106 107 103 105.6

7 103 107 108 10k 107 108 108 107 105 106.3

6 10^ 108 109 101* 109 110 109 108 105 107.1*

5 103 105 109 100 107 109 107 106 103 105.1*

1* 97 96 101* 96 99 103 103 91* 98 98.9

1 91 91 93 87 90 92 95 92 90 91.2

2 87 87 90 87 92 91 90 90 87 89.1

3 81* 86 89 86 90 89 87 89 88 87.6

1* 82 85 88 81* 89 87 86 86 85 85.8 CD 5 81 85 83 82 \J1 86 83 85 83 83.7

6 79 81 83 78 82 82 80 82 79 80.7

7 76 71* 76 77 82 76 80 79 77 77.1*

8 65 69 67 62 75 71 70 69 66 68.2

9 1*8 5o 53 1*7 50 55 53 53 50 51.0

1 121* 132 131* 127 126 125 131 130 - 128.6

2 118 121* 132 123 121* 122 122 121* - 123.6

3 111 120 126 120 119 111* 120 116 - 118.3

1* 106 115 120 113 111* 112 110 113 - 112.9

5 101 108 116 110 109 109 109 109 - 108.9

6 96 105 110 101* 101* 105 101* 101* - 1 0 1 * . 0 27 TABLE 6

Measurements in mm. of female Quiscalus quiscula versicolor wing and tail feathers from Put-in-Bay, Ohio.

Individual Specimens A B C D E F G Average

9 90 9k 91 91 93 90 96 92.1

8 95 97 95 9k 97 92 98 95.1*

7 96 100 96 93 97 92 99 96.1

6 101 I H 96 96 9k 9U 93 99 96.1 3 | 5 92 98 9U 93 88 90 97 93.1 PS Ph 1* 88 93 90 88 82 85 92 88.2

l 82 81* 83 82 82 80 87 82.9

2 80 81 81 78 82 79 8U 80.7

3 79 80 79 76 80 76 82 78.9

1* 77 78 76 76 78 75 78 76.9

5 71* 76 7U 73 78 72 77 71*. 9 r*7•s-l l % 6 72 73 72 69 77 69 76 72.6 si i o o 70 w 7 71 73 69 Ik 6k 67 69.7 CO 8 61 61 72 59 6o kB 61 59.9

9 1*6 1*6 1*6 Ii5 k3 k3 k7 1*5.1

1 111 111 111 106 107 103 111 108.6

2 108 108 108 106 106 102 109 106.7

3 103 108 107 100 101 99 108 103.7

k 98 103 101 99 98 98 103 100.0 3 E-t 5 95 101 98 91* 9k 92 100 96.3

6 91 95 92 90 89 87 9k 91.1 28 TABLE 7

Analysis of variance of Table 5* (Freund et al., 1960j61).

Male Feathers Wing Tail c 1,079,11*7 61*6,352 SSA 1*1*1 585 SSB 32,559 3,1*17 SST 33,118 1*,092 SSE 118 90

Source of Variation Degrees of Sum of Mean Freedom Squares Squares

Between A's (Wing) 8 2*24.1 55.1 Between B's 1U 32,559 2,325.6 Error 112 118 1.1 Total 131* 33,118 -

Between A's (Tail) 7 585 83.6 Between B's 5 3,1*17 683.1* Error 3$ 90 2.6 Total h i M 9 2

*Fa (Wing) - *Fa (Tail) = 32.2 ; F#01 a 3.21 50 • F.01 - 2.67

*Ffi (Wing) r 2,11U i F ,01 = 2‘2^ *Fg (Tail) = 262.8 5 F #01 a 3.60 * = significant

Key

G = correction term SSA a Between A - classifications sum of squares SSB a Between B - classifications sum of squares SST s Total sum of squares SSE = Error sum of squares

A a Feather length variation between specimens. B u Feather length variation between individual feathers. 29

TABLE 8

Analysis of variance of Table 6. (Freund et al., 1960:61).

Female Feathers Wing Tail c 675,01*3 1*2 9, 01*8 SSA 1*30 291 SSB 20,829 1,521* SST 21,790 1,861 SSE 918 1*6

Source of variation Degrees of Sum of Mean Freedom Squares Squares

Between A's (Wing) 6 1*30 71.7 Between B ’s H* 20,829 1,1*87.8 Error 81* 918 10.9 Total 101* 21,790 —

Between A's (Tail) 6 291 1*8.5 Between B's 5 1,521* 301*. 8 Error 30 1*6 1 .5 Total 1*1 1,861

*Fa (Wing) = 6.6 ; F>01 = 3-QU **A (Tail) r 32.3 j F<01 = 3.1*7

*Fb (Wing) a 136.1* i F^q1 = 2.32 *Ffi (Tail) b 203.2 j F #01 - 3.70

* b significant

See Table 7 for a key to terms. Sternal variation

In all measurements a definite size differential between the males and females is noted. A summary of these differences for the three subspecies is given in Tables 2, 3 -An average of all wing feather lengths of the Put-in-Bay, Ohio birds shows the male 11 per oent larger than the female. The male tail feather lengths are 13 per oent longer than the tail feathers of the female. The feather ooloration in the female is much duller than the male with the metallic hues more subdues, some­ times very faint. 31

Ontogenetic variation Ontogenetio variation Is most evident in suooes- sive plumages, particularly in young birds. Qulsoalus qulsoula shows good variation only until the end of the first year. The natal down sparsely oovers the young nestling with feathers of a pale sepia-brown oolor. A post-natal moult begins on the third day of nest life, followed by the acquisition of the Juvenal plumage. This plumage is perfected in 25 days (Eyer, 195*0* The whole plumage is dull clove-brown, the body feathers often very faintly edged with paler brown. The tail is darker with purplish tints and the bill and feet sepia-brown, black when older. A dark brown iris is characteristic of a bird in Juvenal plumage. In late July and August the Prebasio I moult (post- Juvenal) is followed by the Basic I plumage (first winter plumage). The whole plumage is blackish, the head and neck glossed with purple or greenish blue. The wings and tail are iridescent purple to black. Bill and feet are black, the iris light brown to pale Jrellow. There is no Prealternate moult, the Alternate I plumage (first nuptial) is acquired by wear. The Definitive Basic plumage (adult winter) follows a complete prebasio moult 32

(post-nuptial) in late July and August as In the previous year. This graokle is indistinguishable from the Basix I plumage. The Definitive Alternate (adult nuptial) is ac­ quired by wear (Bent, 1958). Wood (19^5) studied the sequence of the Prebasio moult (post-juvenal) in the Purple Graokle (Quiscalus qulsoula stonel) at Harrisburg, Pa. He found the first evldenoe of moulting on 23 July of "the feathers along the edge of the wing". It is assumed he means the primaries. I found the proximal primaries number one, two and three missing as early as 7 July on the Put-in-Bay, Ohio graokles. Wood studied the complete moulting sequenoe and determined that the feather groups moulted in the following order: lesser wing-eoverts, greater ooverts, secondaries, forehead, orown, nape, rump, primary-ooverts, upper tail-ooverts, oheeks, neok, baok, belly, under tail ooverts, scapulars, proximal primaries, breast, ohin, distal remige and the median reotrioes. The Common Graokle has only one complete moult per year and the definitive plumages are acquired after the bird's first year. Adult feather coloration is evident after the Prebasio I moult. Bye oolor generally beoomes lighter as the graokle's body feather oolor beoomes darker. However, one oase of dark brown iris oolor in an adult female Common Graokle is documented by speoimen number 10177 in The Ohio State Museum at Columbus, Ohio 33 Specific and subspeolflo distribution The Common Graokle is distributed through east and central United States, to the east and central portions of south Canada (Figure 1) (A.O.U. Cheok-llst, 1957). Quiscalus qulsoula has been subdivided into three subspecies based upon geographic variation. Two of the subspecies are found breeding only in the United States, one in the united States and Canada. The distribution of Qulsoalus quiscula quiscula" Florida Graokle extends from southeastern Louisiana, southern Mississippi, central western and southeastern Alabama, central Georgia, eastern South Caroline, eastern North Carolina and extreme southeastern Virginia, south to southern Florida (A.O.U. Check-llst, 1957). Qulsoalus qulsoula stonel - Purple Graokle breeds from oentral Louisiana, central and northeastern Missis­ sippi, southern and northeastern Tennessee, eastern Vest Virginia, oentral and northeastern Pennsylvania, oentral southern and southeastern New York, and south­ western Connecticut, south to oentral Alabama, northern Georgia, western South Carolina, east oentral North Carolina and southeastern Virginia (A.O.U. Cheok-list,

1957). 3^

Q. q. quiscula Q. q. versicolor =

Q. £. stonei

Figure 1. North American distribution of the three subspecies of the Common Grackle (Quiscalus quiscula). 35

Qulsoalus qulsoula versioolor. Bronzed Graokle has the most extensive distribution of the three subspecies. It breeds from northeastern British Columbia, oentral southern Mackenzie, oentral Saskatchewan, oentral and northeastern Manitoba, western, oentral, and northeastern Ontario, southern Quebeo, southwestern Newfoundland, and northern Nova Sootla, south along the eastern slope of the Bookies to oentral southern and southeastern Colorado, oentral and southeastern Texas, southwestern Louisiana, western and northern Mississippi, northern Tennessee, Kentuoky, western and oentral West Virginia, Central Pennsylvania, central and oentral eastern New York, northern Connectlout, and southeastern Massachusetts; also on Shelter Island at the eastern end of Long Island, New York. Hybridizes with Qulsoalus qulsoula stonel along the line of junction from southern Louisiana to Massachusetts (A.O.U. Cheok-list, 1957)* 3 6 Recognition of related forms Taxonomio discrimination among the Ioterldae is based upon the external morphology, feather coloration and measurements of the wing, tail, culmen and tarsus. An observer can hare a great degree of confidence in external characteristics for species determination. Knowledge of the locality of the specimen in question is Important in the final determination. Blair et al. (1957) hare summarized the family, generic, and species characteristics so that a concise comparison of United States speoies can be made. Measure­ ments of the species are presented in Table 9. The speoies in this family hare a bill about as long as the head and it is stout. The gonys and depth of the bill at its base is less than the distance from the nostril to the tip of the maxilla. The oulmen is more or less swollen and its tip sometimes deflected. The nostrils are exposed and operculate. There are nine primaries, the outer webs of several are sinuate. The tail is more than half the length of the wing, rounded or graduated, the reotrioes twelve. The tarsus is slightly longer than the middle toe with claw; aorotarsium soutellate; anterior toes slightly ooherent at the base. 37

TABLE 9

Bange of measurements in mm. of United States species in the family Icteridae (Blair et al., 1957).

Species Wing Tail Culmen Tarsus

Tangavius aeneus 98-121; 6i;—98 20-21; 26-32

Molothrus ater 86-116 58-80 ll;-20 23-28

Xanthocephalus xanthocephalus 110-1h6 79-109 20-25 30-37

Agelaius phoeniceus 88-lM 63-106 17-27 25-3U

Agelaius tricolor 105-121; 7i;—95 20-21; 26-31

Euphagus carolinus 103-117 7U-93 18-22 30-32

Euphagus cyanocephalus 116-131; 87-107 19-21; 29-33

Quiscalus quiscula 117-153 101-1U0 28-36 32-39

Cassidix mexicanus 132-201; 118-235 31-U9 37-55

Ictei*us spurius 69-83 6U-75 15-18 21-23

Icterus cucullatus 7U-90 75-99 17-22 20-2U

Icterus graduacauda 90-103 89-106 22-28 25-28

Icterus parisorum 95-106 79-92 21-25 23-26

Icterus galbula 85-102 66-80 16-20 23-26

Icterus bullockii 90-103 70-92 17-21 21;—26

Sturnella magna 89-129 53-87 27-37 35-li6

Sturnella neglecta 105-129 51-83 28-37 3h-h2

Dolichoryx oryzivorus 85-102 59-70 15-18 25-29 The maxlllopalatlnes are weak and curved. It is a New World family with 35 genera and 88 species. In the United States there are 10 genera and 18 species. A summary of the characteristics of Qulscalus is inserted here for comparative purposes. Culmen longer than middle toe with claw, its tip more decurved. The outer primary is shorter than the fourth and the tail much more than 3/4 of the wing, graduated and plicate. The following characteristics of genera in the family Icteridae found In the United States are limited to those features which delimit the genus Qulscalus from them. Bill shorter than head and a rounded tail are characteristics of the genus Tangavlus Lesson. The one United States species Tangavlus aeneus (Wagler), Bed-eyed Cowblrd is much smaller than the Common Grackle

CTable 9). A bill shorter than middle toe without claw and primaries unnotched on Inner web will distinguish viie genus Molothrus Swainson from Q.ulsoalus. The only species Molothrus ater (Boddaert), Brown-headed Cowbird is brown and much smaller than Qulscalus. Genus Xanthooephalus Bonaparte has a tail nearly 3/4 the wing. The only species Xanthooephalus xantho-

cephalus (Bonaparte), Yellow-headed Blackbird has an orange yellow head, neck and breast. 39 A oulmen longer than middle toe without claw and second, third and fourth primaries longest discriminates the genus Agelalus Vielllot. This genus has two United States species. Agelalus phoenloeus (Linnaeus), Red- winged Blackbird and Agelalus tricolor (Audubon), Trioolored Redwing have red lesser coverts. The genus Euphagus Cassln has the outer primary equal In length to the fourth primary. The Rusty Blackbird, Euphagus oarollnus (Muller) has rusty brown upper parts and underparts cinnamon buff. The Brewer's Blackbird, Euphagus oranocephalus (Wagler) breeds west of the Rookies. The genus Cassldlx Lesson has a bill longer than the head and a tall often longer than the wing. There are two species Cassldlx ma.lor (Gmelln), Boat-tailed Grackle end Cassldlx mexloanus (Vielllot), Great-tailed Grackle. The genus Icterus Brisson has a strongly rounded tall. There are six species In the United States. Icterus spurlus (Linnaeus), Orchard Oriole, Icterus ououllatus Swainson, Hooded Oriole, Icterus graduaoauda Lesson, Blaok-headed Oriole, Icterus parlsorum Bonaparte, Scott’s Oriole, Icterus galbula (Linnaeus), Baltimore Oriole and Icterus bullookll (Swalnson), Bullock's Oriole all of which are brightly colored with orange.

A tall less than 3/k of the wing and rounded in the genus Sturnella Vielllot. The two species are 40

Stumella magna (Linnaeus), Eastern Meadowlark and Stumella negleota Audubon, Western Meadowlark, both have a yellow throat, whioh is not present in Qulsoalus. The genus Dollohonyx Swainson has a bill shorter than the head and a straight oulmen. The outer primary is the longest. The one speoies Dollohonyx oryzlvorus (Linnaeus), Bobolink has a golden buff hind neck and is muoh smaller than Qulscalus. Voice

The sounds of the Common Graokle are hard to under­ stand, distinguish and analyze because of their un­ musical patterns. Many authors have likened it to the sound of a rusty hinge. Arlton (19^9) says this of the graokle, "The bird has no song, and there is no music in his harsh conversational ohatterings." Saunders (1935) questions the singing ability of the graokle but admits that it does have a song. Nice (19^3:1^) defines song as follows, "Bird song is a properly sustained, more or less uninterrupted repetition of one or more notes conforming recognizably to a constant specific type and sometimes used by the male as an expression of independent sovereignty." The most obvious sound of the graokle is the "song", a throaty high-pitched squeak accompanied by feather erection and wing spread. It occurs in both male and female, but it is much more vigorous and aggressive sounding in the male. Fioken (1963) calls the male song the "Buff-out Squeak" and the female song the "Buff-out Chuga". This song is associated with the sexual displays before the female, with territorial advertisement and sometimes with aggressive or attack

kl k z behavior. Flcken (1963) describes the "Huff-out Squeak" as follows, "This begins with a simultaneous spreading of the wings and tall and ruffling of the contour feathers ... . As spreading and ruffling reach their maximum, the bird rises up on Its legs, utters the "Squeak", flashes the nlotltans arhythmically, and may take a step or two forward. (The entire display lasts two to four seconds.)" The "Chaok" call of the male and female corresponds to the call note of other passerines and Is the charac­ teristic sound given In flocking situations, either on the ground or In flight, during approaches to the nest or mate, In oases of low Intensity alarm and In solitary birds for no speoial reasons observable to me. under these same ciroumstances a soft, low Intensity, short "Wift, Wlft" is uttered by the male or female, especi­ ally when their mate Is close or approaching. It gives the impression that the bird Is forcing air out of its partially opened bill. Flooking is Induced to some degree when the "Chaok” call note Is given under conditions of alarm. There is a more intense sound to signal alarm which I have divided into the "alarm note" and the "intense alarm". The "alarm note" is given by both the male and female and sounds very much like the Starlings (Sturnus vulgaris) "alarm note", a repeated "Kerr, Kerr, Kerr" with a slightly guttural sound. This will attract a local flock to the scene of the disturbance, the filook generally giving the "Chaok" call. The cause of the "alarm note" was noted to be either the investigator or the gray squirrel (Solurus oarollnensls). The "Intense alarm" was caused by my handling of the nestlings for weighing and measure­ ment purposes. The call is a louder, more rapid sequence of the "alarm note" which attracted grackles into a sizable flock with their "Chack" calls louder and more rapid. Tall flicking aocompanled the two alarm notes, the frequency and intensity increased during the "intense alarm". Tail flicking is an intention movement for flight and is discussed by Andrew (1956). The development of the voice is slow, not reaching full maturity until the following year. On the second day of nest life, a faint "squeak" was heard for the first time. Their voices increased in intensity each day until the eighth day when a change to the "alarm note" occurred. By the eleventh day the young grackles could give the "intense alarm" causing flocking of the adults and attack flights at the investigator while he was hand­ ling the young. The alarm notes and the "Chaok" were the only sound the young could emit until late July when males were seen attempting their "song". Eyer (195*0 44 watched three young captive grackles through the first winter and found that they did not aohieve the full adult song during his study which lasted through the spring* Minor variations in song and in the behavior associated with them are discussed by Floken (19^3)• As discussed by Putnam (1949) voloe probably has a role in species recognition, communication and individual recognition among most birds. Several examples of behavlo* were noted that could be used in support of this thought. During my developmental study at Nest 8-64, the "alarm note" attracted a flock of grackles into the area that certainly must have been alerted to the distress of one of their kind by voice rather than by sight. Some grackles were banded in a shed out of sight, yet their calls attracted a flock of grackles, not other species. The nestlings were seen in the "head up" feeding posture, yet no adults were at the nest, but overhead the "Chaok” call was given by an adult in flight. Communication between pairs seemed especially evident when the "Wift, Wift" call was exohanged during the approach of one mate to another. The feeding cries of the young grackles must stimulate the adults to gather food for them, at least to some extent. During flocking, on the ground or in the air, the persistent *5

"Chaok" must help to maintain the Integrity of the flook. The pre-oopulatory behavior of the male which Includes the "Ruff-out Squeak" and the female** response must be a mutual communication lndloatlng readiness for copu­ lation. Individual recognition by voice must exist to some extent between pairs and between adults and their young. During the first few days of nest life when brooding was quite frequent, the male would approach the nest with food and give the "call note" whereupon the female would fly off the nest from her brooding position. This occurred even If the male approached the female from behind and below out of her sight. There are other factors which probably Induce the female to leave, but voice is certainly one of them. After the young grackles leave their nests, they stay well- concealed In brush or woods, except for their cries for food. Their respective parents seem to have no difficulty finding them, even when other young are numerous in the area. Sound must have a more important role In identification of the young than does sight. Courtship Behavior

Pair formation Pair formation is a lengthy process beginning shortly after the arrival of the females to the breeding grounds, and continuing, in some cases, into the nest building phase. Arrival of the first grackles from the wintering ground occurred in the Columbus, Ohio area on 13 February, 1963, two days earlier than the earliest date recorded by Trautman (1940) for the Buckeye Lake, Ohio area. It is estimated that two days later the grackles appeared at the Put-in-Bay, Ohio area approximately 113 miles north of Columbus. The majority of the females arrived about one week later. The pair bond is achieved through two stages* (1) maintenance of contact between individuals of both sexes and (2) acceptance of the male by the female. The morning hours from dawn to two hours past sunrise are the times most actively devoted to pair behavior, although pairing behavior is evident throughout the day but not with the intensity common in the early morning hours. The trans­ ition between the winter flocking behavior to relative pair isolation dominant during the breeding season is gradual. By the first week of April a few pairs have achieved a stable bond, although the majority are actively engaged in pair formation behavior. Late April and early 46 47 May Is the time for nest construction when most pairs have become stable; however, some pair formation behavior was noticed during nest site selection. The maintenance of a close association between two birds of the opposite sex is the most important aspeot of pair formation. In the early stages, it is probable that a group of males becomes interested in a female and the one which maintains a successful close association against all competition is the eventual mate. This can only be proved through marked birds, large numbers of which are difficult to oapture and mark. The close association of the female with one or more males is evident in feeding, searching for nest materials, resting and especially fly­ ing. Multi-male flights are quite common in the early stages of pair formation. I recorded as many as five males and one female in a flight. Fioken (1963*62) saw as many as six males after a female in flight. In the latter part of April, as pair bonds are becoming established, flights involving one or two males are muoh more common. Most of these flights are of a more moderate flight speed with the female in the lead or the "leader flights" of Fioken (1963*61). Some, however, are chase flights in which the male or males are engaged in high speed pursuit of the female. The male seldom touches the female during chase flights, though bill contact with the female's tail, oausing a rapid direction change in the flight, was noted. The majority of flights occurring during the period of pair bond formation are initiated by the female, with one, two or more males Joining the flight. Most flights originate from a restricted area, that is, a tree, feeding ground, etc., some males, however, Join the flight after it is already in progress. The typical flight, whether a "leader flight" or a chase flight, will find the female in the lead with the male in a direct line to the rear. The female Initiates and terminates the flight, the male following her lead. At flight termination, the male will Invariably seek a higher perch in the same or close tree so that he has a dominant position over her. This behavior changes as the pair bond becomes stronger and is stabilized. Vocalization during flights is discussed by

Fioken (1963:62). The close association between members of the sexes is most evident during late April and May. In early April, feeding is common in groups with a close pairing association being observed in only a few pairs. Most activities throughout the day, after pairing is accomplished, are concerned with the maintenance of the relationship between the sexes. This is especially obvious during feeding sessions. Grackles tend to feed in flocks, yet they maintain their pair integrity in the group. The most common threat display used by the male to help form and maintain the pair bond is the "bill up" display. An Intruding male is approached by a male in the "bill up" position and the distance between them is decreased by a side-stepping movement by the displaying male. The intimidated male will usually retreat, thus in effect increasing the space between males to a tolerable distance. Indication that the female has accepted a male and the pair bond formed is given by variation in two acti­ vities oommon during the pair formation. Obviously, the flights involve only the female and one male with the following variation in the flights* (1) the female does not initiate all flights, the male will fly first, although not as often as the female, and (2) the male no longer strives to achieve a dominant perch at the end of the flight. Keeling of the tail is an important display used by the male during pair formation and paired flights. In a keeled flight, the tail will be strongly V-ed at the beginning of the flight and at the end, the middle por­ tion of the flight is somewhat less extreme, particularly in flights over 100 feet. The degree of keeling at the initiation and termination of flight can be either high or moderate and is sometimes not keeled at all. Keeling is 50 most Intense during the early morning hours and during pair formation and nest building phases. Later In the day and In the breeding season keeling Is less frequent. Data on keeling Intensity and frequency were obtained from 27 April to 3 May, 1965* during the three hours past sunrise. Of 228 paired flights a high degree of keeling was noted In 75 per oent of flights, moderate keeling In 14 per oent and no keeling In 11 per oent.

Sexual behavior The first copulation in the Put-in-Bay, Ohio colony

was noticed on 10 May, 1965* one day after the laying of the first egg in the colony. Pair 10-65 was seen engaging in pre-oopulatory and oopulatory behavior at least once In the early morning and onoe In the early evening during the days that their eggs were laid. Copulation was accomplished In a tree 15 feet from their nest tree and at approximately the same level. As soon as the clutch was oomplete, I did not observe further oopulatory activity. Details of the displays and the copulation sequence are presented by Pioken (1963*65-68). The Nest

Nest anatomy A measurement of four nests yields the following average dimensions: outside diametery 18.4 centimeters; Inside diameter (rim), 10.2 centimeters; outside depth, 15.2 centimeters; Inside depth, 7.8 centimeters; area of nest opening, 82.3 square centimeters. The mean weight of the four nests Is 218 grams. Nest weights varied from 140 to 307 grams, the difference attributed to size £f the nest and dryness of the decayed organic substage• The nest can be conveniently divided Into three parts: the foundation, substage and stage. A detailed breakdown of one nest provided an estimate of kinds of materials used In construction, the number of pleoes and the weight of each of the three major nest divisions. The foundation oontalns the heavy, coarse materials and Is of the simplest construction. There were 271 pleoes used divided among the following: woody stems and leaves - 13* monocot stems - 103» and fine stems - 155. The weight of the foundation was 148 grams. The nest foundation Is Intertwined among the branches or fork supporting the nest to form a cup-shaped bottom and a framework for the sides.

51 The substage conforms to the contour of the cup- shaped stage and gives the nest stability. Included in this part are nest sides and bottom made of stems and grasses solidified with decaying plant matter. There were J26 pieces used in the substage divided among: leaves - 1^, coarse stems - 180, and fine grasses - 132. One source of the decaying plant material was the edge of a eutrophio pond oonneoted to Lake Erie. It is placed at the bottom of the substage oup and provides stability to the nest in ad­ dition to cementing the substage to the stage. The weight of the substage without the deoaying organic matter was 50 grams, the total weight of the substage was 100 grams. The stage was constructed of carefully placed fine grasses and Juniper leaves, which was only a lining to the pre-formed oup of the substage. There were 5^0 pleoes in the stage weighing 8 grams. The total nest weight was 256 grams with 1137 pieces in all three parts.

Location of the nest The Peach Point colony on South Bass Island, Ohio was nested in Juniper (Junlperus vlrglnlana) with the exception of one nest found in boxwood, an ornamental. The lowest nest in the colony of 31 nests was seven feet, the highest nest was 35 feet and the median height was 20 feet. The nest was plaoed in the dense leaves toward the end of a branch or on 53 a main fork, if well concealed. Data accompanying the Dr. B.R. Bales egg collection at The Ohio State Museum gave nest trees and heights for kO nests in Ohio. The number and kind of nest trees with the median neight follow: pine, 26 nests, median height,

25 feet; juniper, 10 nests, median height, Zk feet; apple, 2 nests, median height, 20 feet; elm, 1 nest, height un­ known; osage orange, 1 nest, height, 12 feet. Speolflo names for the trees were not given.

Nest building The graokle builds a well-concealed, deep-oupped nest which is used for one brood for one season. It is difficult to make generalizations about nest building in the oolony because of the striking variation evident from pair to pair. The amount of time spent on nest construction can vary from a few days to over a month depending on pair behavior. The female collects and places all nesting mater­ ials; the male accompanies and guards the female throughout the period of nest construction, however the male was seen carrying nesting materials during the nest site selection. The earliest date for nest construction, which does not inolude the nest site seleotion period, was in late March and the last day devoted to construction in the oolony was

19 May, 1965. It was noted that many more nests are started than are completed and used for egg laying. 54

I had 11 nests under close observation during the nest building phase and only four of them were completed and used. A description of the history of two nests will show the differences between pairs encountered. Pair 10-65 chose the site for their nest on 9 April, on which date the female oarried materials to the site four times between 0655 and 0806 hours.^ The preliminary nest building attempts are evidently part of the nest site selection procedure for they are not part of the true nest building sequence. During the following four weeks, I found little activity at the nest site, only an occasional visit, but without construction materials. Aotual nest construction began on 6 May with six visits between 0605 and 0704 hours or about 10 visits per hour. The foundation was completed on the third day with six visits per hour and an average of 62 seconds devoted to construction each visit. The fourth day was the start of the substage construction and an inactive day with only three visits between 0605 and 0810 hours.

All times were Eastern Standard and a hour clock was used. 55

The fifth day had 33 visits with nesting materials between 0625 and 0910 hours or 12 visits per hour. The average time spent by the female constructing during each visit was 78 seconds. The sixth day was devoted to the decaying plant or "plastering" (Eyer, 195*0 portion of the substage. It was the most active day in the entire building sequence with 40 visits from 0636 to 0905 hours. This averaged about 16 visits per hour and completed the substage construction. The seventh day, 12 May, was the final day of nest building and was confined to the stage or lining. There were seven visits per hour with an average time at the nest each visit of 195 seconds, the longest time spent shaping the nest during its construction. This nest was completed in seven consecutive days with a total number of visits reoorded during the construction of 124. This represents a fairly accurate estimate of construction activity, although some visits were noticed during two afternoons but were not reoorded. The first egg was laid on 15 May, three days after completion of the nest. In contrast to the above orderly building process, Nest 24-65 was constructed in a sporadic fashion, begin­ ning in late March and continuing at Irregular intervals until 19 May. When I arrived at the oolony in early 56

April, the nest had a completed foundation. Work on the nest substage began on 13 April* There were two visits each day with nesting material on 1^ and 15 April, but the materials were dropped. These infrequent visits continued until 21 April then no visits were recorded until 26 April with only one visit occurring on that date. Building on the substage was recorded in the early morning from 7 May through 10 May, with an average of 6.5 visits per hour. Another period of inactivity from 11 May through 17 May was broken by a completion of the substage on 18 May. The stage was completed on 19 May in one day as was the case with Nest IO-65. The first egg was laid in Nest 2^-65 on 22 May, three days after completion of the nest, which

is the same interval recorded for Nests 10-65 and 8-65* Territory

The graokle does establish a limited territory around the nest site, but it does not conform to any of the standard definitions of territory. It comes closest to Nice’s (19^1) definition of the Type D territory usually associated with colonial nesters. Establishment of territory is closely associated with selection of the nest site. The female chooses the nest site and appears to be unrestricted with respect to the area she may investigate. After the site has been chosen, the male will defend that small area in the nest tree which the female has designated as the future nest site. This area defense is limited because the male and female do not remain close to their area until nest build­ ing begins. During nest construction, the strongest territorlalism for the nest site is exhibited by the male. With the advent of egg laying until the young fledge, the female is very closely attached to the nest and male territorial behavior is probably directed to the female rather than the nest site. This strong attraction of the male for the female is evident away from the nest area. The male will Intimidate any challengers for his mate and repel any aggression against her.

57 58

Nest construction can be either very peaceful with few territorial clashes occurring or it can be an almost oonstant fight for the guarding male. The latter usually is the result of two pairs choosing the same nesting area. Vicious fights over this conflict of interest were observed, sometimes nearly to the death for one of the males. The high conflict nests usually were not com­ pleted, the area being abandoned by both pairs. After the female has settled into her nest with sggs, territ-orialism exhibited by the male is probably directed to the female, although the female and the nest have fused into one object. The male responds most vigorously to any interference by the investigator, and to the other extreme usually is unresponsive to other graokles and small songbirds near the nest. There is a great deal of variation from pair to pair in the degree of their responsiveness. Some are more sensitive to in­ trusion than others. For example, Pair 22-65 would permit my investigation of the nest with little complaint. Some­ times the pole-mirror was over the nest before she would fly, the male never responding to this. On the other hand, Pair 2^-65 was very sensitive to my presence. The male faithfully stood guard in a tree 25 feet from the nest and my presence anywhere near the nest tree would elicit an alarm note, which made the female leave the nest. I am certain the graokle is chased by other birds more than it chases. Once the nest is established the male is very tolerant of other grackles near the nest, but it has been observed chasing other species. This also is dependent on individual aggressiveness. I have seen the Cedar Waxwing and House Sparrow steal nesting materials without response from some pairs, but others would give chase to this offense by other species. The grackle will repel the Red-wing if either the male or female come too close (within 8 feet) to its nest. This usually occurs when the two species' nests are close together. The graokles will desert a nest with eggs if disturbed enough, but once the eggs are hatched, the pair become strongly bonded to their young. Any amount of disturbance, short of nest destruction, will not discourage them from maintaining the young. Concomitant with this strong bond is aggressive defense behavior by both male and female. Certainly territorality is involved, but to a greater degree it is an innate bonding to the young. Territorality, then, is strongest during the earliest stages of the breeding cycle, that is, nest site selection and nest construction. During the progressive stages of the cycle, territorality, at least in part, gives way to protection of the female and young by the male, in lieu of a "territory" around the nest. Eggs

The egg The description of the egg was derived from notes made on the Put-in-Bay, Ohio eggs and from a comparison of 28 sets in The Ohio State Museum with the color standards in Palmer (1962). Egg measurements were obtained from three sources. Put-in-Bay, Ohio eggs were measured with a vernier caliper by the author; the measurements of the B.R. Bales Collection in The Ohio State Museum were made by Dr. Bales and found in the notes accompanying the collection; and through a personal communication with F.W. Preston of Butler, Pa., measurements of eggs were obtained from four different areas. Preston (1953) has developed a unique method for measuring and determining shape of bird's eggs. The appearance of the egg shell is dependent upon the intensity and plaoement of the blackish brown pigment over the egg surface. The absence of the dark pigment, although not common, allows the egg's ground color to show through unblemished, a pure pearl gray. At the other extreme the dark pigment can be so heavy, that the ground color is obscured leaving the egg a blotched sepia. The majority of the eggs have a pearl gray ground color with streaks of blaokish brown 60 pigmentation variously located over the surface of the egg. The dark pigment is generally located in greater density at the large end of the egg. The streaking varies from small spots located generally over the surface, to large blotches of dark pigmentation on two or three small areas, or just to limited streaks over part of the egg surface. Using 256 eggs from three sources, I calculated a mean length of 28.14 millimeters, a mean breadth of 20.27 millimeters and a mean elongation of I.38 milli­ meters (Table 10). The shortest length measured was 24.50 millimeters and the longest length 32.00 millimeters. The shortest breadth was 17.50 millimeters and the longest breadth 22.15 millimeters. These average measure­ ments are comparable with Gross in Bent (1958) who gives a mean length of 28,85 and a mean breadth of 21.95 milli­ meters for 20 eggs. Preston, in Palmer (1962) found that time could be saved by measuring one egg, chosen at ran­ dom, from each set instead of measuring all eggs in a set.

He found this to be true for several species, so assumed it would probably be valid for most eggs. Using Bales Collection from The Ohio State Museum and accompanying data I found the mean length for all eggs from 49 sets 62

TABLE 10

Summary of egg measurements from the Dr. B.R. Bales Collection at The Ohio State Museum, F.W. Preston Laboratories and Put-in-Bay, Ohio, eggs.

Bales Bales Preston Put-in-Bay Totals (all eggs) (one/set) and Mean

Length

Total Length (mm) 6,263.00 1,367.75 616.00 32U.30 7,203.30 No. of Eggs 221* k9 21 11 256 Mean Length (mm) 27.96 27.91 29.33 29.50 28.11* Variance 1.72 1.62 1.1*7 U.77 1.80 Standard Deviation 1.31 1.27 1.21 2.18 1.31* Range 21*. 50-32.00

Breadth

Total Breadth (mm) 1*,509.25 938.2$ 1*1*5.78 23i*.60 5,189.63 No. of Eggs 221* k9 21 11 256 Mean Breadth (mm) 20.13 20.16 21.22 21.33 20.27 Variance 0.53 0.58 0.39 0.51 0.52 Standard Deviation 0.73 0.76 0.63 0.71 0.72 Range 17.50-22.15

Elongation

Length/Breadth (mm) 1.38 1.38 1.38 1.38 1.38 63 to be 27.96 millimeters and the mean breadth 20.13 milli­ meters. These figures are not significantly different from the mean length of 27.91 millimeters and mean breadth of 20.16 millimeters obtained by taking only one egg at ran- don (used random numbers table) per set. These measurements are compared in Table 10. The elongation calculation (length/breadth) is the same for all three sources of egg measurements. This shows that all measurements made were consistent in the length and breadth rationand if an error in length or breadth measurements was made, the error was consistent. A t-test (Snedecor 19^6:81) was run on length and breadth measurements to determine if significant differences existed between sources of egg measurement data. Using Preston's measurements as a standard, a comparison of his measurements with Put-in-Bay, Ohio, and the Bales Collection measurements was made. The differences in length and breadth measurements between Preston and Put-in-Bay, Ohio measurements were not significant, but the differences between Preston and the Bales Collection measurements were significantly different (Tables 11 and 12). 64

TABLE 11

Statistical summary of a comparison of mean egg measurements of Put-in-Bay, Ohio, eggs and data from the Preston Laboratories.

Source of No. of Degrees of Mean Sum of Variation Eggs Freedom Length (mm) Squares

Put-in-Bay 11 10 29.50 47.66 Preston 21 20 29.33 29.30

sumrr32 sum=30 diff. of x r 0.17 sum of Sx2 m 76.96

Pooled variance - s r 76.96/30 * 2.565

Standard error - sx -^/s^n^-t- r^J/rqng X ^ 2 . 565(11 + 21)/(11)(21) = 0.60

t = 0 .17/0.60 a 0 .28* 5 d.f. r, 30

Source of No. of Degrees of Mean Sum of Variation Eggs Freedom Breadth (ran) Squares

Put-in-Bay 11 10 21.33 5.09 Preston 21 20 21.22 7.88

suma32 sum=30 diff. Of X r 0.11 sum of Sx2 c 12.97

2 Pooled variance - s - 12.97/30 - 0.432

Standard error - sX = ^ s 2(nL+- x ^ /n ^ p ^ 0.432(1H-21)/(11)(21) « 0.15

t = 0.11/0.15 = 0.73* i d.f. = 30

* a not significant s P 0.5 65

TABLE 12

Statistical summary of a comparison of mean egg measurements of the Dr. B.R. Bales Collection at The Ohio State Museum and from the Preston Laboratories.

Source of No. of Degrees of Mean Sum of Variation Eggs Freedom Length (mm) Squares

Bales 221* 223 27.96 382.91 Preston 21 20 29.33 29.30

sum=2l*5 sum=2l*3 diff. of 5c * 1.37 sum of Sx2 » 1*12.21

Pooled variance - s2 « L12.21/2L3 - I.696

Standard error - sx V s 2(n1 + n 2)/n1n2 J V 1.696(221*-t- 2l)/(22l*)(21) - 0.25 t C 1.37/0.25 = 5.U8** 5 d.f. . 21*3

Source of No. of Degrees of Mean Sum of Variation Eggs Freedom Breadth (mm) Squares

Bales 221* 223 20.13 118.78 Preston 21 20 21.22 7.88

sum=2l*5 sum=2l*3 diff. Of I s 1.09 sum of Sx2 * 126.66

Pooled variance - s2 = 126.66/21*3 - 0.521 Standard error - sx Y s2(nL+ n2)/n1n2 V 0.521(221* + 2l)/(22l*)(2l) - 0.16 t s 1.09/0.16 * 6.81** ; d.f. s 2l*3

■a* = significant : P 0.01. 66

Egg laying Egg laying times were established for 13 nests. Of these four survived to fledge young. Egg laying begins within three days of nest completion. The earli­ est egg was laid on 5 May 64, and the latest egg on 27 May 64, in the Put-in-Bay, Ohio colony. The eggs are laid approximately every 24 hours and all were laid in the early morning except one which was laid in the afternoon. Checks at the nests after the evening obser­ vation revealed no return of the female through the night, and early morning nest cheoks were accomplished before the eggs were laid in six nests (Table 13)• The maximum number of eggs found in a completed set was five and the minimum was three. The factors which control the numbers of eggs are unknown. However, a nest disturbance will cause the graokles to cease laying and desert the nest. Nest 5-64 was disturbed by removing some branches for a better view on 11 May. The female did not desert that same day, but laid one more egg on 12 May and then deserted. This might mean that she is unable to stop the egg laying process immediately, but needs a 24 hour period for cessation. I base this on the fact that if an egg set is complete, the female will desert immediately if disturbed in the manner discussed in the nest success section. Also, she may have an innate 67

TABLE 13

Egg laying times for 13 nests of the Common Grackle (Quiscalus quiscula versicolor) at Put-in-Bay, Ohio.

Nest Egg Date and Time of Date and Time of Number Number Last Observation First Observation Before Laying After Laying

1-6U 1 6 May - 1000 hrs. 6 May - 1900 hrs. 2 6 May - 1900 hrs. 7 May - 1000 hrs. 3 7 May - 1800 hrs. 8 May - 1000 hrs. it 8 May - 1900 hrs. 9 May 0900 hrs. 5 9 May - 1900 hrs. 10 May — 0830 hrs.

2-61t 1 5 May — 1900 hrs. 6 May — 1000 hrs. 2 6 May - 1900 hrs. 7 May - 1000 hrs. 3 7 May - 1800 hrs. 8 May - 1000 hrs. U 8 May - 1900 hrs. 9 May - 0900 hrs. 9 May — 1900 hrs. 10 May — 0830 hrs.

3-6I4. 1 5 May a m 1600 hrs. 6 May rnm 1000 hrs. 2 6 May - 1900 hrs. 7 May - 1000 hrs. 3 7 May - 1800 hrs. 8 May - 1000 hrs. it 8 May — 1800 hrs. 9 May — 0900 hrs.

it-6lt 1 8 May — 1900 hrs. 9 May — 0900 hrs. 2 9 May - 1900 hrs. 10 May - 0830 hrs. 3 10 May - 1900 hrs. 11 May - 0930 hrs. k 11 May - 1925 hrs. 12 May - 0725 hrs. $ 12 May — 1800 hrs. 13 May — 0800 hrs.

$-6k 1 10 May _ 1900 hrs. 11 May — 0930 hrs. 2 11 May — 1930 hrs. 12 May — 0730 hrs.

6-6k 1 8 May _ 1800 hrs. 9 May — 1000 hrs. 2 9 May - 1900 hrs. 10 May - 0900 hrs. 3 10 May - 1915 hrs. 11 May - 0930 hrs. it 11 May - 1930 hrs. 12 May - 0730 hrs. 5 12 May — 1800 hrs. 13 May — 0800 hrs.

20-6U 1 25 May _ 1900 hrs. 26 May — 0830 hrs. 2 26 May - 1800 hrs. 27 May - 0700 hrs. 3 27 May - 1830 hrs. 28 May - 0730 hrs. 68

TABLE 13 (contd.)

Nest Egg Date and Time of Date and Time of Number Number Last Observation First Observation Before Laying After Laying

8-65 1 19 May - 0520 hrs. 19 May - 0620 hrs. 2 20 May - 0530 hrs. 20 May - 0630 hrs. 3 21 May - 05U5 hrs. 21 May - 0635 hrs. k 22 May — 0535 hrs. 22 May - 06U5 hrs.

10-65 1 15 May _ 0530 hrs. 15 May _ 0630 hrs. 2 16 May - o5ii5 hrs. 16 May - 06U5 hrs. 3 17 May — 0550 hrs. 17 May - o6U5 hrs. it 18 May - 06 oo hrs. 18 May - 0715 hrs. 5 19 May — 0615 hrs. 19 May - 1000 hrs.

22-65 1 9 May — 051*5 hrs. 9 May — 0700 hrs. 2 10 May - 0530 hrs. 10 May - 0630 hrs. 3 11 May - 0520 hrs. 11 May - 0620 hrs. it 12 May - 05U5 hrs. 12 May - 0625 hrs. 5 13 May — 0630 hrs. 13 May — 0725 hrs. _ _ 2U-65 l 22 May 0530 hrs. 22 May 06U5 hrs. 2 23 May - 0600 hrs. 23 May - 0715 hrs. 3 2h May — 0530 hrs. 2it May — 0630 hrs.

26—65 1 15 May _ 0530 hrs. 15 May _ 0715 hrs. 2 16 May - 0515 hrs. 16 May - 0650 hrs. 3 17 May - Q5U5 hrs. 17 May - 0700 hrs. It 18 May - 0700 hrs. 18 May — 0800 hrs. U\ cr\ O o 29-65 1 13 May _ hrs. 23 May _ 1100 hrs. 2 m May - 0600 hrs. lii May - 06U5 hrs. 3 15 May - 0530 hrs. 15 May - 0630 hrs. It 16 May - 0530 hrs. 16 May - 0630 hrs. 5 17 May - 05U5 hrs. 17 May - 061i0 hrs. behavioral pattern which directs her to lay in the nest rather than indiscriminately, thus the return for egg laying followed by desertion. During the egg laying period the male was not noticed in the nest area and the nest could be checked for the eggs without any response from the male. However, the female would protest if she were in the hest area. In Nest 9-64 the female spent 25 per oent of 4.5 hours sitting on the nest the day the first egg was laid. On the second day she spent 18 per oent of 7»3 hours sitting on the nest. In Nest 20-64 on the day egg number 3 was laid, thus completing the set, the female incubated 28 per oent of 8.3 hours of observed time. As discussed in Van Tyne and Berger (1959) one must not assume that incubation has started Just because the femalfe is sitting on the nest. It is interesting to note, however, that time is spent on the nest before com­ pletion of the egg set.

Clutch size Clutch size data were derived from a literature survey and from this study at Put-in-Bay, Ohio. Included in the analysis are data from four states and one province, comprising 187 nests and 881 eggs. The mean clutch size is 4.7 which is between the 4.5 given by Eyer (1954) and the 4.9 given by Petersen and Young (1950) (Table 14). 70

TABLE ll;

Clutch size of the Common Grackle (Quiscalus quiscula versicolor) based on incubated or full clutches. a

Area Authority No. Nests No. Eggs in Clutch

Pennsylvania Bales Coll.(0SU Museum) 0 0

Ontario Bales Coll.(0SU Museum) 2 a Bales Coll.(OSU Miseum) 2 0 Bales Coll.(OSU Museum) 1 6 Bales Coll.(OSU Museum) 1 7

Minnesota Bales Coll.(0SU Museum) 1 a

Wisconsin Petersen and Young (1900) 3 3 Petersen and Young (1900) 9 a Petersen and Young (1900) 36 0 Petersen and Young (1900) 6 6 Petersen and Young (1900) 1 7 Buss (1900) 2 a Buss (1900) 0 0

Ohio Bales Coll.(0SU Museum) 2 3 Bales Coll.(OSU Museum) 21 a Bales Coll.(OSU Museum) 18 0 Bales Coll.(OSU Museum) 3 6 OSU Museum 2 3 OSU Museum 13 a OSU Museum 10 0 OSU Museum 2 6 OSU Museum 1 7 Trautman (I9U0) 8 a Trautman (19U0) 8 0 Trautman (19U0) a 6 Trautman (I9U0 ) 1 7 Davie (1898) 2 7 Present Study a 3 Present Stucjy a a Present Stucfy 10 0

Mean a.7

a Trautman's (19U0) figures include clutches and young. Incubation

The female maintains the temperature of the eggs ft throughout the entire incubation period without help from the male. The male role during incubation is restricted to guarding the nest and maintenance of the pair bond by his presence near the nest site during most of the daylight hours. Burns (1915*277) lists the genus Quiscalus as one in which both the male and female take regular turns at incubating. Gross in Bent (1958) and Eyer (195*0 both found the female to be solely responsible for incubation. Certainly, incubation is the responsibility of the female, al­ though the male could be rarely involved. This concept is related to a minor role one male played in brooding, to be discussed later. The factors which control the length of the incuba­ tion period have been discussed by many authors. Incuba­ tion normally begins with the laying of the last egg and ends when the last egg hatches. Temperature is probably the most important factor governing the length of the inoubation period, although humidity is worthy of con­ sideration. Kendeigh (1952) found a high constant temperature maintained by direct sunlight speeded the development of House Wren (Troglodytes aedon) embryos

71 72 by as much as two days shorter than the average Incubation time. Skutch (1962) has discussed in some detail the relationship of the attentiveness of the parents to the length of the incubation period. If a greater percentage of time spent Incubating eggs does hasten the development of the embryo, what might be the survival value? Skutch (1962) answers the question in this way: We might look for such advantages in two directions: constant sitting might increase the safety of the parents or nest, or of both, or it might accelerate the hatching of the eggs. This reduction of the incubation period should less directly reduce losses by diminishing the time the eggs are exposed to predation. A pop­ ulation of birds whose incubation period is 15 days and which loses 30 per oent of its nests to predation before hatching . . . suffers an average loss of 2 per cent per day; so that a reduction of the incubation period to 12 days should increase its hatching success by about o per cent. It has been shown that the incubation period does vary in length, but the effects of the constancy of incubation have been minimal. The constancy of incubation can be calculated from the following equation (Skutch, 1962):

100 S T = ----- S + R

where T = percentage of time on the nest, S = average length of the sessions and R = average I&ngth of the recesses. He believes that a constancy of incubation 73 below 60 per cent will retard hatching, but feels that an increase in constancy over 80 per cent will not shorten the incubation period in passerines. The abnormal incubation period, that is, that period of incubation not terminated by hatching has been little discussed in the literature. Skutch (1962:145) has summarized the articles dealing with this phase of the incubation sequence. In most species the incubation drive does not subside until the incubation time has elapsed one and one-half to two times over the normal period. Berger (1953) found the American Goldfinch (Splnus trlstls) to incubate infertile eggs 23 days or 10 days over the normal 13 day incubation period. Odum (1942) measured a 24 day abnormal incubation period for the Carolina Chickadee (Parus carolinensls) whose normal period is 12 to 13 days. In this study female grackles incubated in one case for 18 days and another for 23 days, or five and ten days over the normal period. Nest 20-64, which had a normal incubation period was compared with Nest 6-64 which did not. The mean percentage of attentiveness or constancy of incubation was only slightly lower in the abnormal nest, 67 per cent in Nest 6-64 and 71 per cent in Nest 20-64 (Figures 2 and 3). There is some tendency toward a Figure 2. Incubation Incubation of Gracklepatterns a Common at female 20-61*Nest 2. Figure duringnormal one theday to day 13 incubation period. Bars represent time on represent timetheneston Bars offtheand time spaces nest.between,the 13incubation day period. DAYS OF INCUBATION 10 12 II 13i 2 3 3 - 6 4 - 5 8 9 - 9 I A 7 -

-

- -

~~r 7 8 H Hh ~9 1 --- —IH 1— 1 lb— I— 11— IIb b 1h H ! — H I — IH IH b HI— 0 I 2 I 12 II 10 H HHHb H ----- OR F DAY OF HOUR H H b H h 1 I lb HI— ---- 1 ---- t— I 1- -I 2 H I—II— IHH H M H HH T I 1 IH H H 1 I H H H l I Ml— H T 6

M

2 2 -

2\-

2 0 - > — 11--- 11— 1

§ 19-\ ^ M HI—IH h H I II---

18- H H I—c >---- —I 17------II--- I---- 1 Ll O !G- I---1

$ is-\ H HF H H I- 14-

13 - H h

12 - -I I-

~ r ~r i —r~ 7 8 9 10 II 12 5 6 HOUR OF DAY Figure 3. Incubation patterns of a female Common Grackle at Nest 6-6U during an extended incubation -o period beyond the normal 13 days. Bars represent time on the nest and the spaces between, time off ^ the nest. Space between horizontal V*s indicate periods of no data. 76 reduction in constancy of incubation during the last three days at the normal and abnormal nest. Also, there was greater variation in attentiveness from day to day in the abnormal nest than was true in the nest with a normal incubation period (Table 15)• Data from four nests studied in 1964 were combined to show variation in the inoubation constancy with respect to time of day and day of incubation. These observations totaled 218 hours, 136 hours of which were spent at nests with incubation periods from day one to day 13 and 82 hours at nests in which the incubation period exceeded the nor­ mal 13 days. (The average normal incubation time to the nearest hour is discussed below.) Tables 16 and 17 pro­ vide a f>er cent attentiveness comparison between data based on day one to day 13 and from day 13 plus. Mean per cent attentiveness based on all 1964 nests with data obtained during day one to day 13 is 73 Per cent. This is in contrast to the 55 per cent mean attentiveness for all data obtained from nests which exceeded day 13 with only the days in excess of the normal incubation period considered. These results indicate that there is some difference in attentiveness between days one to 13 and day 13 plus. In this case the female grackle is 18 per cent more attentive from day one to day 13 than she is during an extended incubation period. TABLE 15

Incubation patterns of 8 female Common Grackles (Quiscalus quiscula versicolor) from Put-in-Bay, 0.

Female No. Day of Hours, No. of Sessions in Recesses in Incubation Minutes, Sessions Minutes Minutes Constancy Temp. Watched Range Average Range Average AM Pi 0700 160<

3-61* 8 2:50 7 li-19 10.6 3-1*2 16.0 1*0 67 70 9 1:00 2 1U-22 18.0 7-17 1 2 .0 60 62 69 10 2:36 6 6-18 12.3 5-27 11.2 52 62 72 11 1:58 6 7-12 9 .6 8-23 1 3 .0 1*2 75 71 12 1*:17 10 3-21* 11*. 0 5-20 10.6 57 50 58 13 1:08 1* 6-11 9.3 6-10 7.7 55 59 63 Hi U:58 11 3-23 9 .7 9-32 17.1* 36 70 81* 15 0:1*0 1 — 20.0 - 2 0 .0 50 75 81 16 5:32 9 3-15 6.1* 17-51* 30.1* 17 75 68 17 1*:00 7 6-16 11.1* 7-55 22.7 33 62 71 18 8:30 2 5-150 75.5 150-305 227.5 26 61* 66 6-61* 7 3:00 1 - 180.0 --— 75 71 8 2:00 1 - 10.0 — 100.0 9 50 58 9 1:25 1 - 85.0 - - - 59 63 10 1*:55 2 25-220 122.5 10- 1*0 25.0 83 70 81* 11 0:55 1 — 3 0 .0 - 25.0 55 75 81 12 5*1*0 1* 50-95 69.5 12-25 1 8 .0 59 75 68 13 1*:25 2 70-11*5 107.5 — 25.0 81 62 71 11* 2:55 3 25-60 38.3 15-25 20.0 66 61* 66 15 7:35 5 20-95 5o.o 15-35 21*.2 67 63 65 16 7:55 2 65-108 86*5 15-270 100.6 1*6 58 61 17 7:05 1* 31-130 83.5 9-37 18.2 82 50 58 18 6:20 5 7-80 33.1: 10-31* 18.6 61* 61 60 19 11:10 10 12-125 1*2.1* 6-27 11*.3 75 58 58 20 3:05 1* 20-50 31.7 7-20 11.6 73 57 61 21 7:15 7 20-70 3 8 .0 10-22 16.5 70 60 62 22 5:1*0 1* 29-65 1*0.2 15-1*8 27.2 60 55 63 23 6:35 7 20-4*5 31*.3 10-1*0 20.6 62 69 60 TABLE 15 (contd.)

Female No. Day of Hours, No. of Sessions in Recesses in Incubation Minutes, Sessions Minutes Minutes Constancy Temp. Watched Range Average Range Average % AM PM 0700 1600

9-61* 1 i*:35 3 10-1*0 21.6 30-85 52.5 29 62 71 2 7:15 1 - 75.0 - 360.0 17 61* 66 20-6h 1 6:00 1* 15-55 31*.5 30-132 71*. 0 32 58 61 2 7:00 5 17-130 60.8 8-15 11.2 81* 50 58 3 5:25 3 12-66 1*6.0 8-29 15.6 75 61 60 1* 11:10 9 23-210 65.2 7-12 1 0 .1* 86 58 58 5 7:17 8 15-9U 1*1*.0 8-15 12.1 78 57 61 6 7:15 5 11-113 5i*.o 11-12 11.2 83 60 62 7 5:55 1* 23 - H 5 82.0 7-10 9.0 91 55 63 8 8:50 7 13-200 67.1 7-12 10.0 87 69 60 9 9:50 8 15-170 61.2 5-30 17.5 78 60 61 10 8:20 10 13-90 31.8 7-20 13.6 70 60 65 11 3:30 5 10-15 23.6 15-30 23.0 51 63 70 12 8:20 8 10-95 1*3.0 10-55 22.1 66 61* 68 13 2:U5 3 20-35 26.6 10-50 28.3 1*8 78 86 22-61* 12 5:26 21 1-25 7.2 1-16 5.8 55 61 60 13 11:05 25 2-60 1l*.l* 2-16 5.7 72 58 58 8-65 2 9:20 11 10-73 38.6 7-20 13.5 71* 56 58 3 10:10 11 15-110 1*3.5 5-25 13.2 77 57 60 1* 9:00 15 9-65 25.1 5-20 10.9 69 65 80 5 7*10 8 10-9I* 38.6 5-35 15.1 72 75 70 6 9*55 5 15-165 105.0 10-30 17.5 86 68 70 7 3:00 2 1*0-130 85.0 - 10.0 89 60 59 8 5:00 5 25-11:0 61*. 8 6-12 9.0 88 50 55 9 11: Olj. 15 10-105 32.7 5-20 12.1* 73 62 59 10 7:1*0 10 15-75 35.8 10-15 11.3 76 68 65 11 10:55 8 25-115 66.2 10-1*0 25.0 72 71* 71 -n3 12 11:00 6 1*2-135 93.7 7-13 9.6 91 61* 55 0 0 13 11:00 8 20-173 71.8 7-15 12.0 85 68 57 TABLE 15 (contd.)

Female No. Day of Hours, No. of Sessions in Recesses in Incubation Minutes, Sessions Minutes Minutes Constancy Temp. Watched Range Average Range Average % AM PM 0700 1600

10-65 1 8:50 15 10-58 23.0 5-20 12.2 63 65 6U 2 12:35 20 10-70 25.7 ii-22 10.0 72 57 60 3 12:35 19 10-70 28.1 5-20 12.1 70 60 67 h 10:30 20 5-U5 20.3 5-17 11.0 65 68 69 5 9:50 12 8-85 38.U 5-20 10.7 78 56 58 6 10:20 lli 5-96 35.6 5-15 9.2 79 57 60 7 9:00 17 k-k$ 21.8 li-20 10.6 67 65 80 8 7:05 8 15-80 Ul.6 5-30 11.5 78 75 70 9 9:5U 15 5-83 3 k .k 5-20 9.1 79 68 70 10 3:00 3 2U-100 53.0 10-11 10.6 83 60 59 11 5:00 h 15-105 50.7 10-17 12.3 80 50 55 12 12:00 9 26-109 57.7 6-35 15.7 78 62 59 13 7:25 10 13-UO 19.3 5-120 25.2 1:3 68 65 2ii-65 2 6:15 11 2-38 16.8 5-28 10.6 61 65 80 3 7:15 12 8-37 18.3 5-70 21.8 U5 75 70 h 10:20 11 5-95 iil.O 9-59 19.9 67 68 70 5 3:00 2 85-85 85.0 - 10.0 89 60 59 6 5:05 3 50-103 67.7 12-15 lii.O 83 50 55 7 12:00 9 15-160 67.6 6-20 12.3 85 62 59 8 7:25 8 5-75 U2.5 10-25 13.1 76 68 65 9 11:15 11 20-105 ii7.2 9-20 lii.2 77 7ii 71 10 9:10 8 15-iiiO 58.2 7-20 10.5 85 6U 55

-o VO 80

TABLE 16

Attentive behavior of Quiscalus quiscula versicolor females8- during the normal incubation period (days 1 through 13) in relation to time of day, at Put-in-Bay, Ohio.

Year 0600-1000 hrs. 1000-11*00 hrs. 11*00-1800 hrs.

1961*

No. of attentive periods 38 77 56 Average length (minutes) 31.7 35.1 39.7 No. of inattentive periods 37 68 52 Average length (minutes) 12.2 11.6 15.1 Total observed time (min.) 1658 3U97 3008 Attentiveness {%) 72 75 72

1965

No. of attentive periods 110 138 97 Average length (minutes) h9.k U7.0 1*6.3 No. of inattentive periods 112 136 78 Average length (minutes) 111. 7 13.0 12.1 Total observed time (min.) U578 6661* 1*1*71 Attentiveness ($) 77 78 79

a 3-6U (part); 6-61* (part); 20—6Uj 22—6U; 8-65; 10-65; 2l*-65. 81

TABLE 17 £L Attentive behavior of Quiscalus quiscula versicolor females during the period beyond the normal incubation period (day 13 plus) in relation to time of day at Put-in-Bay, Ohio.

Year 0600-1000 hrs. 1000-lii00 hrs. Hi00-1800 hrs.

1961*

No. of attentive periods xU 36 31 Average length (minutes) 37.1 2 9 .h 35.5 No. of inattentive periods Hi 39 33 Average length (minutes) Uo.3 29.2 17.0 Total observed time (min.) 108ii 2189 1663 Attentiveness (%) U8 50 68

a 3-6k (part); 6-6k (part). During the first 13 days of incubation, the attentiveness appears to be fairly constant during all the daylight hours with little variation with respect to time of day. In 1964 the per cent attentiveness during the morning hours (0600 - 1000 hrs) was 72 per cent, during midday (1000 - 1400 hrs) 75 per cent and in the afternoon (1400 - 1800 hrs) the per cent atten­ tiveness was 72 per cent. In contrast, there was some variation in incubation attentiveness in relation to time of day during the days past the normal incubation period. The afternoon hours (1400 - 1800 hrs) had the greatest per cent attentiveness with 68 per cent, the morning and midday hours had an approximate 50 per cent attentiveness. Incubation data taken in 1965 included only nests with normal day one to day 13 patterns. The lack of any significant difference between incubation constancy with respect to time of day was found to be

true in 1965 as was 1964. See Table 16 for a comparison of the two years. Observations of sessions and recesses with accom­

panying constancy percentages are presented on a daily basis in Table 15. There is a definite relationship between the air temperature and per cent of incubation constancy by the female. Figure 4 represents 107 hours of incubation observation at Nest 10-65 and shows the o o temperature -! h comstarcy

A-' -80

80- -70

70- -60

$ UJ 60- -50 cl S PERCENT CONSTANCY K so- U 10

~ r ~l------1--- 1----- 1-- 1------1----- 1--- r~ i

for the session and a one minute minimum and 305 minute maximum for the recess. The incubation constancy or the percentage of time the female spends on the nest incubating eggs is 65 per oent, the average for both years. Constancy tends to be lower in nests which do not hatch eggs, so for a more accurate estimate of constancy, the successful nests (nests which hatch eggs, not to be confused with the successful nest in the nest success section which TABLE 18

Summary of data for incubation patterns of the Common Grackle (Quiscalus quiscula versicolor) from Put-in-Bay, Ohio.

Year Hours, No. of Sessions in Recesses in Total Individual 9 Minutes, Sessions Minutes Minutes Constancy Constancy Watched Range Average Range Average % %

1961* 21*3:22 257a 1-220 1*5.3 1-305 35.7 56° 9 3-61* « 31* 9 6-61* = 67 9 9-61* - 19 9 20-61* . 71® 9 22-61* = 67

1965 281* :09 3U7b 2-173 1*5.9 U-120 13.8 77 9 8-65 a* 81® 9 10-65 = 71* 9 21*.—65 s 78

Both 529:31 6Ql* 1-220 1*5.6 1-3Q5 21*.7 65d

^ No. of recesses 1961* s 21*7 No. of recesses 1965 = 3 2 3 ° % constancy for successful nests I96I* = 69 % constancy for successful nests both years = 73 8 Successful nests (eggs that hatch). 86 must fledge at least one bird) were considered separately. In 196^ the constancy was 69 per cent for successful nests only and in 1965 it was 77 per oent. The average constancy for both years considering successful nests only was

73 per cent. 87

Length of the incubation period Nice (195^) defines the incubation period as the elapsed time between the laying of the last egg in a clutch and the hatching of that egg. The mean length of the inoubation period for seven nests in the Put-in-Bay,

Ohio colony was 13 days and ^ hours. The estimated times for laying of the last egg and the hatohlng of the last egg in a clutch were obtained by a look into the nest before and after the eggs were laid or hatched. In 196^, the laying times were obtained by checking the nests the evening before the eggs were laid followed by a subsequent check after the eggs had been laid the next morning. The validity of this method was checked by looking into the nests during the night to see that the female did not return to the nest until morning and in 1965, checks were made into the nest early in the morning before the eggs were laid. In both years, hatching times were established by before and after checks the day the eggs were hatohed. All checks were made at

hour intervals. In Table 19 is a summary of Incubation period data. Three of the nests had a total hatch in each clutch, and four nests had Incomplete hatches. It would seem reason­ able to assume that those nests with complete clutch hatches would give the most accurate length of the inoubation period, 88

TABLE I?

Length of incubation period for the Common Grackle (Quiscalus quiscula versicolor) at Put-in-Bay, Ohio. Data from last egg in a clutch to hatch.

Nest Estimated Time Estimated Time Length of Last Egg Laid Last Egg Hatched Incubation

8-61ia 0700-20 May 0800- 2 June 13 days and 1.0 hr.

lo-6£a O6J4.O-I9 May ll|00— 1 June 13 days and 7.3 hrs.

* 26-65a 0730-18 May 1600-30 May 12 days and 8.5 hrs.

20—61i-b 0615-28 May 071*5-10 June 13 days and 1.5 hrs.

8-65b 0600-22 May 0600- h June 13 days and 0,0 hrs.

22-6$b 0700-13 May 0930-27 May ll* days and 2.5 hrs.

29-65b 06U5-17 May 0600-30 May 12 days and 23.3 hrs.

Mean 13 days and U.O hrs.

a ^ All eggs in clutch hatched. All eggs in clutch did not hatch. for then one could be certain that the last egg laid in the clutch did hatch. However, in all fc r nests with incom­ plete hatches, the calculated length of incubation was longer than the shortest period calculated for one of the nests with complete clutch hatches. It is probable then, that in those nests with incomplete clutch hatches, the last egg which did hatch was indeed the last egg laid in that clutch. On this assumption, all seven nests were combined to give the mean length of incubation stated in the first paragraph. The problem of determining incubation period length is so complex that the working definition of Nice (195*0 will have to be accepted until all factors Involved can be studied. Observations at Nest 9-6*+ revealed some nest sitting on each day of egg laying, but a thermocouple attached to the eggs would be necessary to determine if the brood patch were exposed during these times. It has been shown for several species of adult birds that sitting on eggs does occur without applying heat to them (VanTyne and Berger, 1959)* In most nests, the hatching of all but one egg of a clutch occurs the day before the last egg hatches, thus hatching of a clutch is completed in two days. This would mean that Incubation probably be­ gins the day before the last egg is laid. Eyer (195*+) estimated that Incubation did start in the afternoon of the day before the last egg was laid. He checked this by feeling the eggs to see if they were warm. Jull (1938) found that chicken eggs required a longer incubation period if they are not incubated for a short while after laying. This may be a factor in the com­ pression of the hatching period of a clutch into two days. Care of the Young

This section is concerned with the activities of the adult male and female graokles in relation to brood— ing and feeding of the nestlings and to nest sanitation and maintenance. Two nests at Put-ln-Bay, Ohio were ob­ served for 129 hours which included some time each day of nest life. The nests were in juniper about 15 feet high and next to buildings which served as a blind for the observer. The nestlings in Nest 8-6**- were studied each morning to supply the data for the development of the young section of this paper. My handling of the nestlings did not seem to have an effect on their feed­ ing routine. However, as will be discussed later, there is a significant difference in the brooding and feeding behavior between the two nests studied.

Brooding The most striking facts disclosed by the brooding

data are that the male of Nest 8-64 did brood and the percentage and length of time spent by the female brood­ ing is very different for the two nests. In the main, the female has the primary brooding responsibility. During the first five days of nest life, the female will visit the nest for brooding purposes only in addition to the 91 feeding visits followed by brooding. After five days the brooding only visits decline, brooding being accomplished after feeding. The brooding position is assumed by a lateral back and forth movement of the body over the young, the bill, top of head and tip of the tail visible over the nest edge. Occasionally the female will readjust her position and as the nestlings grow larger will attempt to force them into a different position in the nest cup. Often the brooding session is ended by the presence of the male with food, who gives a soft "ohack" and physically moves to the nest edge forcing the female to reluctantly give up her posi­ tion. The nchackM call note is given by the female also at a majority of arrivals to the nest site. The male of Pair 8-6^ brooded approximately ten minutes each day on the third, fourth and fifth days of nest life (Figure 5)» This unusual behavior was not disclosed at West 20-6^ or by Eyer (195^) In his study of the grackles. Of the five brooding visits by the male, three were visits for brooding purposes only and two were feeding visits followed by brooding. The chance for observational error is slim because of the evident sexual dimorphism in external morphology. My study at two nests gives evidence to the thought that much variation can be expected from pair to pair. + ------t- 3-

I 2 3 4 5 6 7 8 9 10 II 12 DAYS OF NESTLIFE Figure £. Mean brooding sessions per hour for each day of nest life by the Common Grackle at Nest 8—6U. Note unusual brooding behavior of the male on days three, four and five. Equally interesting is the difference in the amount of time spent "brooding through the duration of nest life at Nests 8-64 and 20-64. Figures 6 and 7 show the great difference between Nest 8-64 and 20- 64 with respect to time away from the nest and per cent of brooding by the females at each nest. Brooding ceases after the eighth day of nest life in Nest 8-64 but con­ tinues throughout nest life in Nest 20-64. The brooding curve decreases gradually and uniformly until it is ter­ minated in Nest 8-64, while the brooding curve for Nest 20-64 is quite erratic to the end of nesting. Eyer (1954) reports the female spends six per cent of her time brood­ ing during the last five days of nest life. This compares with zero per cent of the females' time brooding at Nest 8-64 and an average of 23 per cent at Nest 20-64 during the last fives days of nest life. The average percentage of females' time spent brooding during the first five days of nest life is nearly identical for both Nests 8-64 (37 per cent) and 20-64 (38 per cent). Eyer (1954) found the female to spend 50 per cent of her time brooding during the first four days of nest life. Figure 8 reflects the similar brooding behavior between Nests 8-64 and 20=64 during the first five days of nest life and also shows the disparity during the latter half of nest life. The per cent of the females' visits which involves brooding drops to Figure 6. Percent of female Common Grackles' time spent away from the nest, brooding, nest, the from away spent time Grackles' Common female of Percent 6. Figure and in nest maintenance for each day of nest life at Nest 8-6ii. Nest at life nest of day each for maintenance nest in and

PERCENT OF FEMALES' TIME 0" ET 4 6 - 8 NEST 100" 0 2 - 0 5 - 0 8 - 0 6 0 3 - 0 9 10 70- - I brooding 4 7 1 I 12 II 10 9 8 7 6 5 4 3 2 AS FEf LIFE OFNESf DAYS wi fo nest from awaij feeding young nest maintenance and

NEST 2 0 -64

away from vest

UZ SO

C 3 0

brood iv a

feedivq youvqavd vest maivtevavce ------t — 1— i------r 1i i r 5 6 7 8 9 10 II 12 DAYS OF NEST LIFE vo Figure 7» Percent of female Conmon Grackles* time spent away from the nest, brooding, feeding young Os and in nest maintenance for each day of nest life at Nest 20-6U. o-----0 NEST 8 -6 4 +---- +NEST 80-6-4

+---

O 80 -

O Q.

/ 2 3 4 5 6 7 8 9 10 II 12 DAYS OF NEST LIFE Figure 8. Comparison of percentage of female Common Grackle visits with brooding at Nests 8- 6L1 and 20--6h for each day of nest life. 98 the females* visits which involves brooding drops to zero after the eighth day of nest life in Nest 8-6*f, while the female from Nest 20-6*1- has an average of 50 per cent of her visits involving brooding until Day 11. After this time the per cent of her visits involving brooding reduces to 10 per cent.

Feeding Both male and female participate in feeding activity and variation in amount of activity was noted between Nests 8-6*1- and 20-6*1-. In the early days of nest life the male would very often chase the female from her brooding position by his feeding visits. Until the eighth day of nest life in Nest 8-6*1-, most feeding visits are followed by brooding, after this time the visits were always for feeding purposes. The feeding procedure was the same for each visit except for minor variations in position and activity. The male or female arrived in the nest vicinity, paused then approached the nest edge and immediately forced the food into the open mouths and possibly into the esophagus. Some attempt was made to distribute the food to other nestlings, but the most aggressive nestling usually took the food. The duration of the feeding act was timed and found to last, on the average, about one minute. The variation in duration 99 seemed to be connected, at least in part, by the type of food. A b i l l full of m a y f l i e s took longer to deposit than did one caterpillar. No variation in feeding tech­ niques between male or female adults was observed. Variation in the role of the sexes at both nests was noted. Data were compiled to show these variations and. are presented in graphic form. Figure 9 compares the total feeding activity at the two nests and shows that the rate of feeding at each nest was similar. During the first five days of nest life an average combined male and female feeding rate of 1.03 feedings per nestling per hour was obtained for Nest 8-64 and 1.16 feedings per nestling per hour for Nest 20-64. Similar results were found for the last five days of nest life when total male and female

feeding activity was compared at these two nests. A feed­ ing rate of 2.55 feedings per nestling per hour was found for Nest 8-64 and 2.10 feedings per nestling per hour for Nest 20-64. Thus at these two nests the combined feeding activity by both male and female adults is comparable. Male and female feeding activity at each of the two nests was very different when the activity of one sex was compared with the other. Figures 10 and 11 show that at Nest 8-64 the male was slightly more active than o NEST 8 -6 4 j . + + N EST2 0 - 6 4

> h

DAYS OF NEST LIFE 100 Figure 9. Comparison of the total number of feedings per nestling per hour by both male and female adult Conmon Grackles at Nests 8-6U and 20-6U for each day of nest life. 3-

+ — + cf

■ + -

DAYS OF NEST LIFE

Figure 10. Comparison of the number of feedings per nestling per hour made by the adult male and 101 female Comnon Grackles at Nest 8-6U. +----- + o ’

— k

DAYS OF NEST LIFE 102 Figure 11. Comparison of the number of feedings per nestling per hour made by the adult male and female Common Grackles at Nest 20-6U. 103 the female, whereas in contrast, the male of Nest 20-64 was very inactive, the female accomplishing most of the feeding tasks. An average of the feedings per nestling per hour for the total nest life time was computed to show the differences in the feeding behavior between the sexes at each nest. The male from Nest 8-64 averaged 1.00 feedings per nestling per hour, slightly more than the females* 0.82 feedings per nestling per hour rate. In sharp contrast, the male from Nest 20-64 had a 0.32 feed­ ings per nestling per hour rate and his mate 1.34 feedings per nestling per hour, considerably more active. In both nests the male started feeding on the second or third day of nest life, leaving the female with the total feeding responsibility early in nest life. During the majority of the nest life period, the male keeps a fairly stable feed­ ing rate until the final days when male activity drops off, gradually in Nest 8-64, suddenly in Nest 20-64. The female response to the male feeding activity decrease was one of moderate activity decrease in Nest 8-64 to one of rapid feeding inorease in Nest 20-64. Very few general­ ities can be made about the feeding activity roles of the sexes because they are so variable from pair to pair. However, a combined male and female feeding activity pic­ ture compares favorably from nest to nest. Feeding activity in relation to time of day is summarized in Table 20. There are no dramatic peak feeding times, although a tendency to increased activity is noted between 0800 and 1000 hours, between 1200 and 1400 hours, and toward the end of the day from 1700 hours until dark. Feeding activity, however, is sustained throughout the daylight hours with little variation.

Nest maintenance This section is concerned with adult activity in relation to nest sanitation and repair. The removal of fecal sacs and probably parasites from the nest cup are the factors involved in nest sanitation. Nest re­ pair concerns the physical remaking of the nest lining and to some extent the foundation. The male and female shared the fecal sac removal duties, but the female was almost completely responsible for parasite removal and

nest repair. A greater percentage of time was spent by the female at Nest 8-64 in nest maintenance than was the case at Nest 20-64. Refer to Figures 6 and 7. The time spent in feeding the young would be nearly the same at both nests, so the larger percentage In the graph reflects, in the main, the Increased maintenance activity at Nest 8-64. The reason for the Increased maintenance at Nest 8-64 is probably due to the tearing of the nest materials TABLE 20

Number of feeding visits by male and female Common Grackles (Quiscalus quiscula versicolor) at Nests 8-6it and 20-6it in relation to time of day.

Hour of Day

6 7 8 9 10 11 12 13 lii 15 16 17 18

Nest 8-6it (5 nestlings)

Days of Nest Life

First four days - - 5 it 6 - 5 it 5 it 7 11

Middle four days - 6 8 5 8 8 10 8 7 10 8 10 10

Last four days —— 11 10 12 9 11 12 10 11 — 10

Nest 2 0 — 61|. (3 nestlings)

Days of Nest Life

First four days 2 3 — 5 3 1 it 5 3 2 It -

Middle four days it 5 7 3 6 5 5 8 3 6 5 5 7

Last four days 7 5 6 7 8 — — 8 7 8 8 9 it

H o 106 as I lifted the nestlings twice a day throughout nest life for the development study. The grasping of the nestlings* feet became stronger later In nest life, and Figure 6 reflects a greater percentage of the females* time devoted to nest maintenance toward the end of nest- llfe. Fecal sacs were emitted by the nestlings at more than half the visits by the adults. Figures 12 and 13 show the percentage of male and female visits during which fecal sacs were emitted by the nestlings and then eaten by the adults at both Nest 8-64 and 20-64. The fecal sacs were eaten in a greater percentage during the early days of nest life, with a decrease In frequency later. The gap between the fecal sac emission line and the fecal sac eaten line represents those fecal sacs carried away in the adult’s bill. The reasons for the Increase in the adults flying with the fecal sacs, as the nestlings become older, might be a decreased need for concealment as the young birds beoome stronger, for the eating of the fecal sac might render the adult less conspicuous. Also the increase in the size of the fecal mass, plus a weakening of the fecal sac wall as the nestlings age might produce a less acceptable food package for the adults. NEST 8-6 A o-----o fecal sac emission

100 - +----+fecal sacs eaten

60- O f

2 0 -

0-

DAYS OF NEST LIFE Figure 12. Nest sanitation by adult male and female Common Grackles at Nest 8-6h for each day of nest life. The gap between the solid and dotted lines indicates the percentage of visits by the adults after which the fecal sacs were carried away in their bill, rather than eaten. NEST 20-64 o----o fecal sac emission

100 - -1----+fecal sacs eaten

80-

20 - ki O-

1 2 3 4 5 6 7 8 9 10 II 12 DAYS OF NEST LIFE 108 Figure 13, Nest sanitation by adult male and female Common Grackles at Nest 20-61* for each day of nest life. The gap between the solid and dotted lines indicates the percentage of visits by the adults after which the fecal sacs were carried away in their bill, rather than eaten. 109

The proportion of fecal sacs carried away from the nest or eaten by the male as compared with the female is fairly equal in Nest 8-64, but the female from Nest 20-64 removed more fecal sacs than the male. Consider­ ing the total nest life period of 12 days, the 8-64 female disposed of 45 per cent of the fecal sacs and the 20-64 female 82 per cent. The males were more active during the last five days of nest life than they were during the first five days. The male at Nest 8-64 carried away 50 per cent of the fecal sacs during the first five days, leaving 49 per cent for the female. In contrast, the male at Nest 20-64 carried away 4 per cent, leaving the bulk for the female. During the last five days of nest life, the male carried away 62 per cent of the fecal sacs from Nest

8-64 and 25 per cent from Nest 20-64. The differences between the activity of the sexes at the two nests are fairly stable regardless of the type activity. The relative inactivity of the male at Nest 20-64 is reflected in feeding, brooding’ and nest maintenance behavior. The reasons for this obvious in­ dividual variation in behavior are unknown. Pledging and Post-nesting Activities

The process of fledging and one day’s activities out of the nest was observed in Nest 20-64. The handling of the young in Nest 8-64 forced them to leave the nest one day earlier than those in Nest 20-64. The young from both nests were color banded but were not seen after the second day past fledging. This restricts the major part of the post-nesting activities to generalities. The first departure from the nest occurred the after­ noon of the 13th day of nest life. The other two nest­ lings left the nest the morning of the 14th day. The method of departure is awkward, in one case the young graokle fell out of the nest and grasped a branch one foot below the nest while another nestling Jumped out of the nest to a branch and immediately took a position near the other fledgling. The adults* attention was divided between those young in and out of the nest until all had fledged. In an attempt to reach a bill full of food, the fledglings fell several times to lower branches, until all fell to the ground, I noticed that the first grasp­ ing attempts by new fledglings were not normal, in that the front (3rd) toe was behind the branch and the hind (4th) toe in front. This condition was corrected about

110 Ill four hours after fledging. Feeding calls became more persistent in fledged young. While nestlings, the calls are given usually when the adult arrives with food, whereas after fledging, the young call continuously from their perch. For two days past fledging the feeding rate re­ mains similar to those calculated for nestlings. It is probable the feeding rate remains constant for at least a week past fledging, although my observations are soant. Total dependency on the adult for feeding appears to last approximately two weeks past fledging. The next two weeks are a learning period for the young although they still are dependent upon the adult. The following period of two weeks is one of semi-independence with independence following six to eight weeks after fledging. During the training period the immature grackles look much like the adult female, so the feeding exchanges are quite noticeable and unusual looking. Feeding activities were watched at a food dump for two weeks following fledging. The immatures remained close to the adults on the ground and in flight. Nearly every adult had an immature grackle in pursuit flight, especi­ ally in the dump area. Toward the end of the fourth week past fledging, some immatures would fly into the dump and forage with the adults, while others appeared to be inde­ pendent in their searches. 112

Flocking became apparent in late July and early August, increasing in intensity through August. Indi­ vidual birds or pairs were noticed less frequently as the summer passed. Foraging over grape vineyards, corn fields or the city dump were commonly noted activities late in the summer. Observations of the roost disclosed flock movements from the mainland about five miles to the south. Wave after wave of blackbirds were seen flying across the water and they were followed at dusk by car to the roost. The grackles in company with Red-wings and Starlings could have started a reverse migration or they could have been local shifts in population. Details on flock movements are difficult to obtain. Nest Success

There are four factors which limit nest successs (1) infertile eggs, (2) weather, (3) predators and (4) human interference. The latter factor would be more pronounced in an observed colony than in a colony under natural conditions. The Put-in-Bay colony studied in 1964 and 1965 contained 31 nests, seven of which were inaccessible, thus limited to binocular observation. In 1964, seven of 15 nests fledged at least one young bird

for a 47 per cent nesting success and in 19^5* three of nine nests were successful for a 33 per oent nesting success. The average nesting success for both years was 4-1 per oent. In this study of nest success, a nest was considered success­ ful if at least one nestling fledged. Pledging means a de­ parture from the nest by the young bird at the end of its nestling period of development. Petersen and Young (1950) In their grackle study in Wisconsin gave a nesting sucoess of 55 per cent over a three year period. Eyer (1964) found a 35 per cent nest­ ing success in his Michigan study over a two year period. The average nesting success for all three areas, Wisconsin, Michigan and Ohio, which Included 119 active nests, was 4? per oent. Nioe (1957) found a nesting success of 46

113 114 per oent In 29 studies of open nest altricial birds. The data presented here fall close to the average found by Nice. The 65 per cent fledgling success based on eggs hatched for all Put-in-Bay, Ohio nests is the same as was found in Wisconsin but is 25 per cent higher than the Michigan study. The number of fledglings produced from eggs laid is a much lower 30 per cent due to the high rate of infertility at the beginning of the nesting season. The causes of fledging failure were weather and starva­ tion in the nest. Other factors could contribute to the loss of fledglings, but these two were the only ones en­ countered in this study (Table 21). Nest failures were common to nests started early in May at the beginning of the breeding season. In 1964, all eight failures in the Put-in-Bay, Ohio colony were nests in which the first egg was laid by 8 May. In 1965* the one failure due to infertile eggs was an early nest, the first egg laid on 9 May. The causes were distributed between infertile eggs, predators, weather and human interference. Weather was the most destructive natural factor due in part to the high position of the nest in juniper and to the high velocity of north-east winds which frequent the Lake Erie islands (Table 22). 115

TABLE 21

Nesting success of the Common Grackle (Quiscalus quiscula versicolor) from Put-in-Bay, Ohio.

T otals and 196U 1965 Averages

Active nests 15 9 2h Percent successful U7 33 hi Number of eggs 60 32 92 Number hatching 2h 15 39 Percent hatching ha hi h2 Number of fledglings 18 8 26 Percent of young fledging 75 53 65 Percent of eggs producing fledglings 30 25 28

TABLE 22

Causes of nest failure of the Common Grackle (Quiscalus quiscula v e r sic o lo r ) from Put-in-Bay, Ohio*

V)6h 1965 Total Percent

Human interference 3 2 5 36 Weather 1 3 U 28 Infertile eggs 2 1 3 22 Predators 2 0 2 Hi 116

The predators most likely to induce harm are the gray squirrel (Solurus carollnensls), Norway rat (Hattus norveglous). oat (Felis famllarlus). crow (Corvus brachyrhynohos) and Blue Jay (Cyanocltta cristata). A gray squirrel was seen taking eggs from a deserted nest and is suspected of killing an incuba­ ting female found torn to shreds beneath her nest. Nest loss due to human interference is caused through a lack of understanding what the investigator can or cannot do in the nest area. The birds are more prone to desert in the early stages of nesting and become strongly bonded to the nest after the eggs hatch. The incubating female will tolerate looking into the nest with a pole mirror, but she will desert if any of the surrounding branches are moved to a different position. One nest was lost be­ cause of cutting some branches above the nest while eggs were in the nest. Another nest with eggs was also deserted when I tied a branch so that it moved six inches away from the nest. After hatching the adults will tolerate branch removal as well as removal of the nestlings for short periods. Adults show the strongest defensive displays during the nestling phase of the nesting cycle. Development of Young

The young In Nest 8-64- were weighed twice, the bill, tarsus and wing measured once a day from hatching until fledging, 1 June to 13 June, 1964. These data are sup­ plemented by daily observations of morphology and behavior. A detailed feather tract study was made on a 12 day old nestling. Feather tract nomenclature follows that of

VanTyne and Berger (1939)*

First day The newly hatohed young were partially covered by natal down in the morning of the first day of nest life and by evening the down had increased in length, so that it covered moderately thickly the back, sides, wings and neck. The skin was pink and soft with the dispropor­ tionately large abdomen the dominant feature. The eyes were closed and remained closed until the fourth day. The head was "all mouth" and could be lifted into gape position for two to three seconds. The bill was very soft, light- colored and seemed to surround the pink and black pattern of the oral membranes, so obvious while gaping. The first removal of a fecal sac by the adult female was noted at 1000 hrs. Photographs of the nestlings were taken for the 12 days of nest life and help show these features in Fig­ ures 14 through 25. 117 118

Figure lU. One day old Common Grackle nestling from Nest 8-6U on 1 June 6U. Weight at 0800 hours was 7.5 grams. 119

Figure If? . Two day old Common Grackle nestling from Nest 8-61* on 2 Jure 6k Weight at 0800 hours was 11,9 grams. Figure 16. Three day old Common Grackle nestling from Nest 8-6U on 3 June 6It, Weight at 0800 hours was 18.U grams. Figure 17. Four day old Common Grackle nestling from Nest8- 6 ii on u June 6U. Weight at 0800 hours was 28.2 grams. Figure 18. Five day old Common Grackle nestling from Nest8- 6 I4. on $ June 6I4 . Weight at 0800 hours was 35.1 grams. 123

Figure 19. Six day old Common Grackle nestling from Nest 8-6U on 6 June 6U. Weight at 0800 hours was U3.2 grains. Figure 20. Seven day old Common Grackle nestling from Nest 8-6U 7 June 6iu Weight at 0800 hours was 55.0 grams. Figure 21. Eight day old Common Grackle nestling from Nest 8—6U on 8 June 61;. Weight at 0800 hours was 58.3 grams. Figure 22. Nine day old Common Grackle nestling from Nest 8-61; on 9 June 6h. Weight at 0800 hours was 6£.8 grams. Figure 23. Ten day old Common Grackle nestling from Nest 8-61* on 10 Jnne 61*. Weight at 0800 hours was 70.3 grams. 128

Figure 2k, Eleven day old Common Grackle nestling from Nest 8-6b on 11 June 6h, Weight at 0800 hours was 75.0 grams* 129

Figure 25. Twelve day old Common Grackle nestling from Nest 8-61* on 12 June 61*. Weight at 0800 hours was 76.2 grams. 130

Second, day The young nestlings doubled their birth weight by evening giving some indication of feeding success. The skin coloration darkened especially on the head, neck, wings and spinal areas. A faintly audible squeak accom­ panied the gaping and an attempt at crawling with their stubby wings and feet was noted. Distinct feather sheaths were seen on the feather tracts but were particularly evident in the primary, secondary, spinal and rectrioe regions. There was little change in behavior, the major activities being gaping and defecation. The neck re­ mained very weak, allowing but short gaping attempts.

Third day In addition to the increase in weight and length, which will be treated later, the developing feather tracts were now quite evident. The primary feather sheaths rup­ tured the skin but were of dark color with no "feather­ ing out" occurring.until the seventh day. Natal down was absent on the ventral surface from birth and the ventral tract became visible. The rectrioes penetrated the skin but did not rupture. Disturbance of the nest­ lings would elicit the gaping response accompanied by a more audible cry. Defecation occurred when the nestlings were removed from the nest and vigorous crawling made weighing difficult. 131

Fourth day The sub-dermal feathers on the capital tract remained the slowest to emerge, the humerals were well-developed and easy to see. The spinal tract had Its best growth In the cervical region, decreasing in length posteriorly to the pelvic region. The primaries and secondaries were plainly visible, and the sheaths of the coverts were noticeable under the skin. Posteriorly, feather develop­ ment was slower with the crural and femoral tracts darkened but not erupted. The rectrlces remained the dominant feather group in this area. The largest nestling, with eyes opened slightly, was able to maintain a position in the nest superior to the others. All nestlings could gape and crawl actively. Some grasping reaction was noted as attempts were made to cling to the nest material. Gaping and defecating remained the principle activities.

Fifth day Little change was noted in the capital, spinal, femoral and crural tracts. The primaries and secondaries continued to develop rapidly, extending well beyond the trailing edge of the wing. The marginal coverts had penetrated the skin with the alulal feathers visible. Ventrally the feathers began penetration of the skin, the interramal, submalar and malar regions form a W-shape 132 from the base of the lower bill to the neck. The remain­ ing ventral tract forms an inverted-Y containing the newly- emerged sternal, axillar and abdominal regions. Increased strength was evident In the wings and legs as crawling attempts made measurement and weighing quite difficult. Removal from the nest triggered the grasping response which was now capable of some nest tearing. A correlation between fecal sac ejection and handling was noted as fecal sacs were left on the balance, the white paper used as a background for the photographs, and even on the camera.

Sixth da.y The changes In external appearance were minimal, however the eyes were at least partially open In all the nestlings. The rectrlces increased their growth rate and were now plainly visible. A slight emergence of the anal circlet and crural feathers was noted. The natal down has been almost completely replaced by the emerging juvenal feathers.

Seventh day The feather shafts in all tracts appeared to have made great growth gains. The very long primary and secondary feather shafts ruptured so that approximately one millimeter of feather was exposed. The capital tract and anal circlet feathers were the last to rupture, so 133 that now a final growth surge would feather the nestling

In three to four days. Eyes were completely open In all nestlings. They were very active and would try to orawl as soon as they were placed on the ground. The feeding response was very strong and would react to a movement of the nest or their surroundings. I could not detect any fear in their behavior, but they appeared to show caution in their movements.

Eighth dav The rapid growth which started on the seventh day continued. Some feathers in all tract areas had ruptured, the primaries and secondaries lengthened the most and the feathers on the occiput the least. The bill had be­ come harder and darker tiwh the conical shape more apparent. Their cries were much stronger and sounded more ma­ ture. Gaping and defecating were still the primary be­ havior. Attempts at standing in the nest while feeding and elevation of the rump for defecating over the nest rim were observed. The five nestlings were color banded, and the smallest one had become feeble.

Ninth Day Little change in behavior or morphology from the previous day. The feathers continued to grow beyond the feather tracts, yet they covered only the tract areas. 13^ A heavy infestation of nest mites was noticed for the first time. The smallest nestling died between 0800 hrs. and 1800 hrs.

Tenth day The feathers were extended well beyond the tips of the sheaths in most tract areas giving a "feathered out" appearance over approximately 80 per cent of the body. All nestlings were able to sit upright and move in all directions on the ground. The grasping strength in the feet was strong enough to support the young grackle on my finger for a short period. Preening attempts and wing exercise were noted in the nest. Some avoidance reactions while handling them indicated the fear response was increasing.

Eleventh day The continued Increase in feathering was most strik­ ing in appearance this day. The body was completely oovered with feathers giving the impression that feather growth was complete. Disintegration of the feather sheaths coupled with rapid feather growth freed the vanes so that the barbs and barbules had spread to their normal position. Removal of the young grackles from their nest became increasingly difficult due to their ability to give the alarm note, which caused attacks by adults on the investi­ gator, and a sudden development of the fear response, caus­ ing a tighter grasping response in the nestling*s feet for 135 nest materials. A decline in vigor was noted in the yellow- banded nestling at the 1800 hrs. check.

Twelfth day Feather growth continued at a rapid rate with all body areas covered except the temporal apterium where natal down was still visible. The nestlings became quite excited when handled, gave intense alarm calls, struggled to escape on the ground and took short flights refusing to re-enter the nest. The yellow-banded nestling died during the day from what appeared to be starvation evidenced by a weight loss started the ninth day.

Thirteenth day All remaining nestlings had left the nest, the white- banded one remaining in the nest tree. The others were unable to be found in the area. The next day the white- banded nestling left the tree and despite intensive searching, none of the fledglings was seen again. This dispersive behavior made a detailed study of them im­ possible. The young grackles remained dependent upon the adults until about the middle of July. They were seen flying in pursuit of the adults, calling in an immature voice, but they were of the same size. Their feather coloration re­ mained brownish, lacking the metallic sheen which is characteristic of the adults, until toward the end of 1 3 $ August, It was difficult to tell immatures from the adults in the field, however in the hand determinations were easier; hut in August it became increasingly dif­ ficult to tell immatures from adult females. Figure 26 shows the rapid increase of weight from hatching to fledging. The lower line reflects the growth progress of the weakest nestling which died during the seventh day of nest life. The nestlings that survived showed a continuing rapid weight increase until fledging. During the tenth through twelfth days some decline in growth activity is reflected in the age-weight curve and in a striking overnight weight loss. The average hatch weight for five nestlings was 6.8 grams with a low of 5.8 grams and a high of 7.5 grams. The average weight of three fledglings (last day of nest life) was 73*3 grams, with a 61.0 grams low and an 81.6 grams high. Daily weight gains were subject to wide fluctuation, in some cases the nestling would gain 10 grams a day and in one case on the seventh day of nest life a loss of 1.7 grams was recorded. It was the only day a loss was recorded and this day was the only day rain occurred during the nest life study. The daily weight gains averaged 5.0^ grams over the 12 day period, which indicates that tremendous growth was occurring in those nestlings aggressive enough to gain their share of the food intake (Table 23). Lack of WEIGHT IN GRAMS 100 20 Age-weight relationshipsin a brood of young Common Graeklesfrom Nest M .PM AM 1 2 3 u =E5z IUE 26 FIGURE AGE IN DAYS AGE 6

7 8 9 8-61*. 10 "S ^ 11 12 H 138

TABLE 23

Weights in grams of Quiscalus quiscula versicolor nestlings from Nest 8-61*, Put-in-Bay, Ohio.

Date Leg Band Color Time Red Blue White Yellow Red-Red

6/ 1/61* 0800 hrs. 7.5* 7.2a 6.5a 6.9 1800 hrs. 10.1 8.8 8.0 9.0 — 6/ 2/6h 0800 hrs. 11.9 10.3 9.3 10.0 5.8* 1800 hrs. 16.8 ll*.i* 11.2. 13.8 7.3 6/ 3/61* 0800 hrs. 18.U 15.7 12.0 11*. U 8.2 1800 hrs. 26.6 20.3 18.1 19.7 10.1* 6/ 1*/61* 0800 hrs. 28.2 22.8 19.2 23.1 12.0 1800 hrs. 32.5 26.5 21*.3 26.0 15.7 6/ 5/61* 0800 hrs. 35.1 29.5 26.6 29.8 17.0 1800 hrs. 1*3.0 33.6 32.1* 31*.3 19.0 6/ 6/61* 0800 hrs. 1*3.2 35.0 33.1* 35.0 20.9 1800 hrs. 53.1 11.0 39.1* 1*0.0 22.1 6/ 7/61* 0800 hrs. 55.0 U2.6 1*1*.o 39.3 21.1* 1800 hrs. 58.0 1*2.5 1*2.3 1*2.2 22.5 6/ 8/61* 0800 hrs. 58.3 1*5.7 1*2.3 37.7 20.5 1800 hrs. 61*.5 1*9.5 50.7 1*3.5 20.1* 6/ 9/61* 0800 hrs. 65.8 5o.o 1*7.7 1*0.7 18.2 1800 hrs. 71.5 55.7 55.6 1*5.6 died 6/10/61* 0800 hrs. 70.3 52.9 51*.6 39.5 1800 hrs. 78.3 57.1* 61*.o 1*2.6 - 6/11/61* 0800 hrs. 75.0 55.6 62.0 38.1* 1800 hrs. 80.8 60.1 66.0 38.8 - 6/12/61* 0800 hrs. 76.2 55.9h 60.9, 3U.1* 1800 hrs. 81.6b 61.0b 68.1*b died

a hatch b fledge 139 aggressiveness would result in less food, which in turn weakened their attempt at food grasping, eventually leading to their death. Once inferiority among nest mates is es­ tablished, it tends to lead to their destruction. Measurements of the bill, tarsus, wing and ninth pri­ mary were made daily, during each day of nest life, on the five nestlings in Nest 8-64- from Put-in-Bay, Ohio. The measurements were made with calipers, the body parts measured in the following manner: bill - along the culmen, from tip to base at skull; tarsus - from middle Joint (behind) between tibiotarsus and tarsometatarsus to end of tarsus in front (to lower edge of last undivided scute); wing - measured from flesh at bend of folded wing to tip of longest primary (to fleshy tip before primaries are exposed); ninth primary - from tip to point of eruption at base of sheath. Measurement procedure is from Palmer (1962:5). Figure 27 shows the average growth curves for four nestlings in Nest 8-64. The typical sigmoid curve is evident for the tarsus and wing growth. A logarithmic representation of these two growth curves yields a straight line indicating the geometric nature of wing and tarsus growth. The bill grows very slowly, increasing slightly the first eight days of nest life, then levels off to the LENGTH IN MILLIMETERS 3° 35 Uo 15 20 10 25 0 Figure 27. The average growth ofgrowth averageGrackle The nestlings four Common 8-6h. Nest from 27.Figure 1 2 3 h DAIS OF NEST LIFE NEST OF DAIS 6 7 8 9 10 11 12 ::::Tarsus Wing Bill Ninth Primary

Otfl 141 day of fledging. In contrast, the ninth primary breaks through its sheath on the third day of nest life than grows very rapidly an almost equal amount each day of the nest life (Tables 24 and 25). No correlation be­ tween growth rates and temperatures in Table 25 could be established. An average body part growth rate of four nestlings was calculated and confirms the visual pic­ ture established by Figure 27. The tarsus and wing grew very nearly the same rate during the nestling period, 2.45 mm/day and 2.43 mm/day, respectively. The bill grew at the slowest rate of 0.68 mm/day and the ninth primary the most rapid at 3-00 mm/day. A study of the pterylosls of a twelve day old nest­ ling from Nest 8-64 and weighing 3^ grams, involved a mapping of the feather tracts along with a feather count in each tract (Figures 28, 29 and JO),

Capital tract This pteryla begins as a narrow band at the base of the bill, widens at a point posterior to the eyes to cover the head to an imaginary line drawn from the angle of the Jaw to the mandibular ramus, then to the base of the skull. There were 553 feathers in this tract. 142

TABLE 24

Length in mm. of the bill, tarsus, wing, and ninth primary of Quiscalus quiscula versicolor nestlings from Nest 8-64, Put-in-Bay, Ohio.

Leg Band Days of Nest Life Color and Body Part la 2 3 4 5 6 7 8 9 10 11 12

Red Bill 8 9 10 11 12 13 14 14 16 16 16 16 Tarsus 13 16 17 23 26 31 35 37 40 40 41 42 Wing 11 13 15 21 24 28 33 33 36 37 38 41 9th primary — — 1 3 5 11 14 19 23 23 32 37

Blue Bill 7 8 9 10 11 12 13 14 15 15 15 15 Tarsus 13 15 17 21 25 30 32 35 37 37 38 38 Wing 10 12 15 18 22 26 28 31 33 34 37 37 9th primary — — — 2 4 10 14 20 25 28 31 34

White Bill 8 9 10 11 12 12 13 14 15 15 15 15 Tarsus 11 12 15 19 24 29 32 35 36 38 41 41 Wing 9 10 12 17 20 26 29 32 34 35 36 36 9th primary — — 1 3 7 10 16 18 24 26 30

Yellow3 Bill 7 8 9 10 11 12 13 14 14 14 14 14 Tarsus 12 15 17 21 25 29 32 35 35 35 35 36 Wing 10 12 14 19 22 26 29 30 31 31 32 33 9th primary — — — 1 4 9 13 16 20 21 25 31

Red-Red*3 Bill 7 8 9 10 10 11 12 13 - - -- Tarsus 11 12 16 18 23 25 27 27 - ——— Wing 10 11 13 15 19 22 24 24 - - —— 9th primary - - - 1 4 7 9 11 - - - -

a 1 June, 1964 b died TABLE 25>

Growth rates in mm. per day of Quiscalus quiscula versicolor nestlings for each day of nest life from Nest 8-6Z*, Put-in-Bay, Ohio.

Leg Band Days of Nest Life Color and Body Part Ia 2 3 5 6 7 8 9 10 11

Red Bill 1 1 1 1 1 1 0 2 0 0 0 Tarsus 3 1 6 ■ 3 h 2 3 0 1 1 Wing 2 2 6 3 h $ 0 3 1 1 3 9th primary 0 1 2 2 6 3 5 h 3 U 5

Blue Bill 1 1 1 1 1 1 l 1 0 0 0 Tarsus 2 2 h 3 2 3 2 0 1 0 Wing 2 3 3 li h 2 3 2 l 3 0 9th primary 0 0 2 2 6 h 6 5 3 3 3

■White Bill 1 l 1 1 0 1 1 i 0 0 0 Tarsus 1 3 h 5 5 3 3 1 2 3 0 Wing 1 2 5 3 6 3 3 2 1 1 0 9th primary 0 0 1 2 h 3 3 3 6 2 U

Yellow*3 SiTl 1 1 l 1 1 1 l 0 0 0 0 Tarsus 3 2 h k h 3 3 0 0 0 1 Wing 2 2 5 3 k 3 1 1 0 1 1 9th primary 0 0 l 3 5 h 3 h 1 h 6

Red-Redb Bill 1 1 1 0 1 1 1 - - - — Tarsus 1 h 2 2 2 0 —-—— Wing 1 2 2 k 3 2 0 - -- - 9th primary 0 0 1 3 3 2 2 m m

f" 1 June, 19&h b died 144

Figure 28. Ventral view of a twelve day old Common Grackle nestling showing the major feather tracts. marginal coverts

alula

ventral tract

primaries

crural tract

Tect rices 146

Figure 29. Dorsal view of a twelve day old Common Grackle nestling showing the major feather tracts. humeral tract

marginal coverts ) capital tract

alula

spinal tract ! primaries

secondaries ?

femoral tract crural tract Iff 1

rectrices H 148

Figure 30. Lateral view of a twelve day old Common Grackle nestlii^ showing the .major feather tracts. capital tract

rec trices

humeral tract

femoral tract ventral tract 150 Spinal tract This feather tract extends from the base of the skull (end of capital tract) to the upper tall coverts. It Is a narrow band of feathers throughout Its length with some widening In the neck and pelvic areas. It is subdivided into four regions; the cervical region extends from the capital tract to the trunk and is bordered on each side by an area free of feathers, the cervical apterium; the interscapular region extends posteriorly between the scapulae; the dorsal region extends from the scapulae to a point halfway to the tail and the pelvic region lying between the pelvic girdle extends from the dorsal region to the upper tail coverts. These last three regions are bounded laterally by the lateral apterium. There were 266 feathers in this tract.

Humeral tract This small pteryla runs obliquely posterior on the dorsal surface of the wing near the wing-trunk junction. This feather tract contains the scapulars whioh sore relatively dense. There were 116 feathers in this tract.

Femoral tract This tract runs along the junction of the leg and trunk, parallel to the spinal tract and extends poster­ iorly almost to the anus. There were 200 feathers counted in this tract. 151 Crural traot The feathers of the leg are In the orural tract. The distribution of feathers is most dense on the ventral aspect of the leg, dorsally the leg feathers are sparse, but increase in density on the distal portion of the leg. There were 244 feathers in this tract.

Alar traot The wing feathers are subdivided into these major feather types: the primaries, their coverts; the secondaries, their coverts; the alula and the marginal coverts. The alar tract includes all the wing feathers except the hu­ meral tract feathers. The feather count of both wings gave a total of 808 feathers.

Caudal traot This tract Includes the reotrices, the upper and under tail coverts and the anal circlet. Dorsally the caudal tract is bounded by the uropygial gland which separates the spinal and caudal tracts. Ventrally the anal circlet ends the tract. There were 94 feathers in this tract.

Ventral traot This is the largest single tract on the grackle. The tract begins in the soft area between the lower mandibles, extends laterally to an Imaginary line beneath the eye, forms a single narrow band in the 152 cervical region, then divercates around the protruding lower neck area and remains as two separate narrow bands running posteriorly along the ventro-lateral edge of the trunk to the anal area. The tract is bounded by cer­ vical and lateral apterla. The central ventral area sur­ rounded by the two lateral branches of the tract is the median apterium. This tract had 103^ feathers. The total number of feathers in all tracts was 3315» slightly higher than the count made by Wetmore (1936) on the Purple Grackle male in 1933 in which he counted 2730 feathers. A male might have a greater number of feathers since he is slightly larger than the female. The sex of the nestling on which my study was based was unknown. Migration

Spring and autumn Factors which Influence the annual population shifts north and south are a complex combination of climate, physiology and behavior. Differences In departure and arrival dates from year to year can usually be traced to climatic differences at the origin and along the migra­ tion route. The spring migration of the Common Grackle begins in the south-central portion of the United States in early February and reaches its climax in the prfcirie provinces of Canada in May. Eyer (195*+s25) has mapped the migratory path giving dates at localities as the wave ad­ vances north. The median date of arrival at the Buckeye

Lake, Ohio area is given by Trautman (19*K)) as 2k February. Eyer (195*0 gives the dates, 25 February to 5 March for Ohio. Maurice Giltz (pers. comm.) reported the first migratory wave at Columbus, Ohio on 13 February, 19^5» with the breeding population settled in by 27 February. Post-breeding dispersal is difficult to study because the methods are limited. Banding provides the only feasible method to date and its value is restricted because of the

153 154 limited time for recoveries. There Is some Indication that at least some grackles fly north from their breed­ ing territory before going south for the winter. Of the 67 recoveries of Ohio banded grackles, taken from the files of the U.S. Pish and Wildlife Service, seven were recovered in August to October. Of these seven, four were recovered in Ontario in September and October. These recoveries were banded in April and July in northern Ohio, which indicates that they were probably on their breeding grounds when banded. If these birds were to fly south again, these data represent a reverse migration of from 75 to ^-00 miles into Ontario. It is probable only a small proportion of the population reverse migrates, if it occurs at all. The flocked grackles leave central Ohio around 20 November (Trautman, 19^0) and arrive on their wintering grounds in late November or early December. 155 Winter range Small numbers of the Common Grackle are found in the northern portions of their range during the winter, although the majority of the wintering populations are located in the lower Mississippi valley and the south­ eastern United States. Monroe (1961) lists Norfolk County, Virginia as the area with the largest concentra­ tion of 5*000,000 grackles found during the 6lst Christmas bird count in December, i960. Monroe (1962) again lists Norfolk County, Virginia, with 7,000,000 Common Grackles counted in December, 1961, as the highest concentration in the United States. Martin (1963) listed Rome, Georgia, with 2,310,000 grackles, for the Christmas bird count in December, 1962, as the largest concentration. Monroe and Gauthreaux (196*0, in their summary of highest counts for the Christmas count of December, 1963* listed Little Rock, Arkansas, with 24,000,000 Common Grackles. The winter range extremes include Halifax, Nova Scotia; Fredericton, New Brunswick; Quebec, Quebec; Ottawa, Ontario; Toronto, Ontario; Saskatoon, Saskatche­ wan; Detroit, Michigan; Appleton, Wisconsin; St. Paul, Minnesota; Lower Souris National Wildlife Refuge, North Dakota; Cheyenne, Wyoming; Longmont, Colorado; and Ft. Worth, Texas. This represents the fringe of the scattered winter range. See Table 26 for summary of sightings. 156

TABLE 26

Number of Common Grackles counted in a 15 mile diameter circle in December for four years at the extremes of its winter range. (Audubon Field Notes, 1961,62,63,6ii)

Locality Year

I960 1961 1962 1963

Halifax, Nova Scotia 15 1 0

Fredericton, New Brunswick 3

Quebec, Quebec

Ottawa, Ontario 1 3

Toronto, Ontario 2 h

Saskatoon, Saskatchewan 0 2

Detroit, Michigan 1

Appleton, Wisconsin 1 3

St. Paul, Minnesota 6 7 5

Lower Souris Nat. Wildlife Refuge, North Dakota 1 0 3 9

Cheyenne, Wyoming 0 5 0

Longmont, Colorado 1 0 0

Ft. Worth, Texas 13 0 75 157 The largest concentrations are found in northeast Texas; northwest Louisiana; southwest Arkansas; north­ west Mississippi; east central Alabama; central and northwest Georgia; southeast, southwest and northwest Tennessee; west central Kentucky; southeastern Ohio; southwest South Carolina; north central North Carolina; southeast Virginia; eastern Maryland; and southeast New Jersey. See Table 27 for numbers and location of sightings. Cook (1888) noted the absence of the grackle over large sections of Texas in winter. Peterson (i960) has outlined the winter range in Texas, which is gen­ erally restricted to east Texas. Eyer (195*0 points out that there is a winter population in northern Mexioo. Some Common Grackles have been spotted along the Rio Grande River to Del Rio, which could be the edge of the Mexican population. Florida does not have a large wintering pop­ ulation, although they are present in varying numbers throughout the state. No large aggregates over 10,000 were found on Christmas counts reported from Florida. Areas with concentrations over 10,000 are found no further north than Cape May, New Jersey, along the Atlantic coast. In the Middle Atlantic States, Buckeye Lake, Ohio has the most northern large concentrations of grackles. (Audubon Field Notes, I96I, 62, 63, 6*0 158

TABLE 27

Highest concentrations of Common Grackles over 10,000 individuals counted in a 15 mile diameter circle in December over a four year period. (Audubon Field Notes, 1961,62,63 ,6k)

Locality Tear No. of Individuals

Tyler, Texas 1963 22,959 Shreveport, Louisiana i960 37,700 El Dorado, Arkansas 1962 1, 775,000 Little Rock, Arkansas 1963 2U,000,000 Texarkana, Arkansas 1961 77,117 Moon Lake, Mississippi I960 2,000,000 Auburn, Alabama 1962 270,000 Mille dgeville, Georgia 1962 1,250,000 Rome, Georgia 1962 2, 310,000 Chattanooga, Tennessee I960 15,578 Memphis, Tennessee I960 900,000 Reelfoot Lake, Tennessee 1961 700,000 Henderson, Kentucky I96I 60,000 Buckeye Lake, Ohio I96I 19,685 Clemson, South Carolina I960 1,000,000 Aiken, South Carolina 1962 77,655 Greensboro, North Carolina 1963 1,800,000 Back Bay Nat. Wildlife Refuge, Va. 1962 23,675 Norfolk County, Virginia 1961 7,000,000 Ocean City, Maryland I960 i,3 k o ,o o o St. Michaels, Maryland I960 250,353 Rehoboth, Delaware i960 500,000 Cape May, New Jersey i960 50,000 159 An indication of the principle wintering areas for Ohio grackles can be obtained from banding recovery reports. The original bandings were made in 19 Ohio counties from 1923 to 1955. These reports with their recoveries were supplied by the Migratory Bird Population Station, U.S. Pish and Wildlife Service, Laurel, Maryland. These data are presented in Table 28 and discussed below. Recoveries during the four month period - November through February - were reported from ten states which include Alabama, Arkansas, Florida, Georgia, Indiana, Kentucky, Louisiana, Mississippi, Tennessee, and Texas. This list agrees closely with a report from Michigan by Eyer (195*0 and an Indiana report by Perkins (1932). The grackles banded in Michigan were not reoovered in Florida or Georgia and the Indiana grackles were not recovered in Ohio or Texas, but they were reported from Iowa. An estimation of the percentage of Ohio grackles in the various states from November through February can be determined from the recovery reports. Of the *1-8 winter return reports, 44 per cent were reported from Tennessee, 23 per cent from Alabama and 15 per oent from Mississippi.

The remaining states had about 2.5 per cent each. Eyer (1954) listed Tennessee with 32 per cent of the recovery returns and Mississippi with 20 per cent. Alabama ranked fourth 160

TABLE 28

Number of reports of recovered Ohio banded Common Grackles (Quiscalus guiscula versicolor) by area and season. Data from the files of the U.S. Fish and Wildlife Service.

State or Season Recaptured Total Province Nov.-Feb, Mar.-July Aug.-Oct.

Alabama 11 1 0 12 Arkansas 1 0 0 1 Florida 1 1 0 2 Georgia 1 0 0 1 Indiana 1 1 1 3 Kentucky 2 1 0 3 Louisiana 2 0 0 2 Mississippi 7 1 0 8 Ohio 0 6 2 8 Ontario 0 0 k k Tennessee 21 1 0 22 Texas 1 0. 0 1

Totals U8 12 7 67 l6l with less than one per cent. Tennessee extends far enough east and west to absorb the greatest share of Ohio and Michigan migrant grackles. The main thrust of migrants heads in a south-south-west direction, so Mississippi gets the next greatest share of Michigan's grackles and Alabama gets the next greatest share of Ohio's grackles. The average distance traveled by the autumn migrants is about 450 miles. The greatest distance was to Texas, an 860 mile trip.

Longevity In Bent (1958:404) there is a report of a recovery of a 17 year old banded grackle, which appears to be the longevity record for the Common Grackle (£.£. versicolor). The oldest grackles recovered from Ohio banded birds are 10 years old. Of the 67 grackle recovery reports, the following numbers of the various ages were founds 10 years old, 3; 9 years old, 1; 8 years old, 3» 7 years old, 1; 6 years old, 2; 5 years old, 1} 4 years old, 3» 2 years old, 10; 1 year old, 40. The mean age calculated from

these reports is 2.5 years. Pood Habits

Introduction The destructive food habits of the grackle are well documented by government pamphlets, articles and testimony by the farmer. Data gathered at Put-in-Bay, Ohio has been incorporated with data from the literature to give a better general summary of the grackle's food habits. In spite of all the adverse criticism found about this bird, it should be said in all fairness that the grackle is in more ways beneficial than harmful. The reader should peruse this section with an open mind and arrive at his own conclusions. Data on food habits have been available in the literature since the latter part of the last century. Changes in farming methods and habitat with the con­ comitant increase of the blackbird population, tend to make less reliable this older data. Nevertheless, it is useful to establish a relationship between past and present trends in food patterns for the Common Grackle.

Adult food types My data and that available in the literature lead one to the conclusion that the grackle is omnivorous 162 and eats what is available at the time. Of course some preference is shown, especially when food is plentiful. The omnivorous food habits have been quite strongly developed in the Icteridae. Beecher (1951) found a strong, well-developed mandible adductor muscle in the grackle which has the same role functionally as the adductor muscle mass in the finch-like Cowbird. In ad­ dition to this muscle, Wetmore (1919) described a hard keel projecting downward from the h o m y palate of the Bronzed and Purple Graokles. This keel is used in a sawing adaptation, which enables the graokles to out through acorns, corn and other hard foods. The evolu­ tionary implications are striking. It is likely Cassidii, which has a plentiful food supply year around without having to crack hard foods, gave rise to the northern Qulscalus which had to be able to make use of hard foods in order to survive. Graokles are continually trying new feeding methods which selection pressure could perfect possibly providing a new advantage to the grackle in his changing environment. An examination of 24 stomaohs from Put-in-Bay, Ohio graokles in July, 1964, consisted of a determination of food types and the dry weights of animal and plant parts. The animal and plant foods were broken into pieces and 164 were in different stages of digestion, making Identification difficult. I was able to classify the plants to genus and the animals to order, in most cases. The foods were separated into animal and plant portions, dried for 12 hours and weighed (Table 29). The calculated mean weights of the birds by sex and the animal and plant food weights fall into an expected pattern. The male grackle's mean weight was 119.2 grams, the female's 95*1 grams and the immature grackle's mean weight was 96.1 grams. The plant food made up the major part of the diet of all graokles examined. The males consumed a mean dry weight for plants of 0.3576 grams, the females 0.2924 grams and the immatures 0.2830 grams. These figures correspond closely to the body weights given above, in that the larger males eat the greatest amount; the females and immatures weighing nearly the same con­ sumed an almost equal dry weight of plant food. With re­ spect to the animal matter and stones consumed, the males again consumed the greatest weight. The immatures, however, had a greater weight of animal parts and stones than the female. A consideration of all sexes and ages together shows a mean dry weight for plant remains of 93*2 per cent, animal remains 4.3 per cent and stones plus other miscellan­ eous items 2.5 per cent. The food types found in the examined stomachs are listed in order of frequency en­ countered: Moras sp, 22 stomachs, Coleoptera. 14 stomachs, 165 TABLE 29

Weight in grains and types of stomach contents from adult and immature Quiscalus quiscula versicolor from Put-in-Bay, Ohio.

Sex Date Weight of Dry Weight Food Type Bird Animal Plant Misc. cf 7 / 7/61* 1U0 0 .01*60 0.3818 Coleoptera Diptera Rubus sp. Morus sp. cf 7/ 8/61* 106 0.01*32 0.3568 Coleoptera Rubus sp. Morus sp. cf 7/10/61* 123 0 .0301 * 0.1*971 Coleoptera Morus sp. cf 7/13/61* 112 0.2000 Solarium sp. Morus sp. cf 7/13/61* 121 0.0082 0.371*2 0.1393 Coleoptera Morus sp. cf 7/13/61* 116 0.0187 0.7255 Coleoptera Diptera Hymenoptera Solarium sp. Morus sp. cf 7/15/61* 125 0.0127 0.1925 Coleoptera Morus ap. cf 7/15/61* 117 0 .1*321* Morus sp. Solanum sp. cf 7/20/61* 113 0.0095 0.0586 0.0300 Araneida Rubus sp. Morus sp.

9 7/ 8/61* 107 0.0012 0.1*1*27 Coleoptera Rubus sp. Morus sp.

9 7/ 8/61* 92 0.0101 0.1903 Oligochaeta Coleoptera Diptera Morus sp. Rubus sp. 166

TABLE 29 (contd.)

Sex Date Weight of Dry Weight Food Type Bird Animal P lan t Misc.

9 7/ 10M 111 O.OO98 0.3197 - Morus sp.

9 7 /1 3 M 88 0.01i;0 0.2890 - C oleoptera Morus sp.

9 7 /1 5 M 88 - 0.27U0 0.0200 Rubus sp. Morus sp.

9 7/15/61; 90 0.0027 0 . 1;700 Rubus sp. Morus sp. Malus sp.

9 7/20/6U 90 0.0161; 0.0611 Oligochaeta C oleoptera Rubus sp.

Imm 7 / 8/6U 95 0.00l;6 0.7372 - C oleoptera Morus sp.

Imm 7/13/6U 115 0.0170 0.3660 - Rubus sp.

Imm 7/15/61; 81 0.0087 0.3031; 0.0793 C oleoptera Solanum sp. Rubus sp. Morus sp.

Imm 7/15/61; 95 0.0052 0.0958 - Morus sp.

Inrni 7/15/61; 103 0.0212 0.1861; - C oleoptera Morus sp. Imm 7/1S/6U 86 0.0186 0.2l;26 — Morus sp.

Imm 7/15/61; 86 0.0192 0.2U51; - Morus sp.

Imm7/15/61; 108 0.0305 0.0869 — C oleoptera Morus sp. 16?

Rubus sp, 10 stomachs; Solatium sp., 4 stomaohs; Diptera, 3 stomachs; Oligochaeta, 2 stomachs; and Araneida, Hymenoptera and Malus sp., 1 stomach each. Beal (1900) made an examination of 2,3^6 stomachs collected during all months of the year. Comparison of his data with that presented above is Impossible because it is not stated what the percentages were based upon.

My percentages were based on dry weights, and I suspect Beal's were based on volumes. Nonetheless his work is valuable and adds to the knowledge of the grackle's food habits. He found a total of 69.7 per cent plant food and 30.3 per cent animal food in the 2,3^6 stomachs, which included ^56 nestlings. Table 30 gives the percentage of food types by months. Determination of plant and insect types taken from the stomachs is presented in Table Jl. Despite the fact that nearly one-third of the food consumed consists of harmful insects, attention is given to their harmful consumption of grains and fruits, which accounts for one-half of their total food Intake. Beal (1900) points out that the greatest amount of corn is eaten in February, which is mostly waste grain and of no value to the farmer. The amount of consumption de­ creases until July, when it reaches the minimum, then in September and October c o m constitutes 53*2 per cent 168 TABLE 30

Food of the Common Grackle. Number of stomachs examined: January, 7; February, 8; March, 53; Ajbril, 289; May, 3U8; June, 887; July, 31:6; August, 197; September, 81; October, 111; November, 11; December, 8. Total 2,3U6. From Beal (1900).

Months Food Ave. J F M A M J J A s 0 N D % % % % % % % % % % % % % (Animal) Pred. beetles 5.6 7.1 0.3 U.3 6.3 8.5 12.9 11.1 9.1 3.1 2.6 1.5 0.5 May-beetles U.9 - 0.1 3.2 6.8 22.0 15.1 8.0 0.8 0.6 0.5 - 1.6 Snout-beetles 2.0 2.9 0.3 1.2 1.8 3.5 5.3 u.u 2.1 l.U o.U 0.1 0.7 Other beetles 1.0 2.3 - 1.2 1.3 1.7 1.9 1.3 0.9 0.5 0.1 - 0.6 Caterpillars 2.3 U.3 0.3 1.2 2.3 8.2 3.6 1.2 0.7 1.3 1.3 - 3.1 Grasshoppers 7.3 - 0.6 0.8 1.2 8.8 1U.6 12.7 23.U 8.1 6.9 9.0 1.9 Other insects 2.3 3.0 0.3 3.8 2.9 3.2 2.9 U.8 1.9 1.7 2.3 0.5 o.U Spiders and myriapods 1.5 1.0 o.5 0.6 2.3 7.U 2.9 1.3 0.5 o.U 0.6 0.5 0.6 Crustaceans and mollusks 3.1 11.7 0.1 1.6 1.9 2.U 0.9 0.2 2.6 2.3 9.9 3.1 Vertebrates 0.3 - - 1.2 0.3 0.7 0.5 0.2 0.1 0.6 Tr. - Total 30.3 32.3 2.U 17.6 26.8 65.9 62.1 U5.9 39.7 20.3 17.0 21.5 12.5

(Plant) Corn 37.2 l.U 82.0 58.U U0.927.2 28.2 7.7 1U.0 53.2 51.5 35.U U7.3 Oats 2.9 - 1.2 0.9 1U.5 2.2 o.U 5.2 9.3 1.1 0.8 - - Wheat U.8 --- 1.2 0.6 0.2 26.1 25.9' 0.8 - - - Other grain 1.6 6.7 - 0.6 0.3 - 0.2 - Tr. 0.8 10.5 - Domestic fruit 2.9 o.U Tr. 5.8 10.3 9.1 1.3 o.U - 8.2 Wild fruit 2.1 -— 1.7 1.2 o.U 1.3 2.7 1.3 1.5 2.2 U.5 8.1 Weed seed U.2 - 7.3 10.6 6.2 o.U Tr. 0.6 0.3 2.5 11.6 11.6 - Nuts 1U.0 66.3 o.U 10.U 8.6 3.0 2.0 1.3 O.U 10.U 15.7 16.5 23.9 Total 69.7 67.797.6 82. U 73.23U.1 37.9 5U.1 60.3 79.883.0 78.5 87.5

Tr. s trace TABLE 31

Identified insects from the stomachs of Common Grackles. Beal (1900).

COLEOPTERA

Cicindela punctulata Bolbocerus farctus Cicindela purpurea Geotrupes sp. Cychrus sp. Dichelonycha elongata Carabus sp. Macrodac tylus~subspinosus Calosoma scrutator bachnosterna sp. Calosoma calidum Anomala varians Calosoma externum Ligyrus gibbosus Pasimachus depressus Allorhina nitida Scarites subterraneus Euphoria fulgida Amara sp. Euphoria inda Chlaenius sp. Cremastochilus sp. Agonoderus pallipes Prionus sp. Harpalus~caliginosus Strangalis sp. Harpalus pennsylvanicus Cryptocephalus venustus Helophorus inquinatus Typophorus canellus Olopnrum convexum Colaspis brunnea Scymnus sp. Chrysomela pulchra Hister americanus Castro idea polygoiii Ips quadriguttatus Haltica sp. Drasterius elegans Dibolia sp. Drasterius dorsalis Coptocycla signifera Podabrus reguTo'sus Eleodes tricostata Canthon sp. Bpicaerus imbricatus Phanaeus carnifex Phytonomus punctatus Onthophagus hecate Sitones hxspidulus Onthophagus pe nnsylvanicus Lixus sp. Ataenius sp. Balaninus sp. Aphodius fimetarius Sphenophorus zeae Aphodius inquinatus Calandra granaria

LEPIDOPTERA

Leucania unipuncta Deilephila lineata

HEMIPTERA

Euschistus sp. Prionidus cristatus

NEUROPTERA

Corydalis cornutus 170 and 51*5 per cent, respectively, of the food totals and this represents damage to the standing corn. Fruit damage is minimized "by Beal's study and only raspberries and blackberries accounted for the 10 per cent fruit found in the grackles' stomachs. Other fruits were found in trace amounts. In areas of high fruit concentra­ tion, such as was found in my study area at Put-in-Bay, Ohio, damage to the grape farmer is economically disastrous unless protective devices sire used. No grackle stomachs were analyzed in September, when grapes ripen, but informa­ tion from farmers forms the basis for this discussion. The grapes raised on the Bass Islands are used primarily for wine manufacture. It was pointed out to me that grackles and Red-wings damage only one or two grapes in a bunch, which spoil and thus in turn yield the whole bunch useless for wine processing. Protective measures are necessary to prevent complete destruction of the crop. The most commonly used devices are the carbide gun, which fires automatically every few minutes, and the shotgun. During the period of peak ripening, personal patrols with the shotgun are necessary in the morning and evening. Mr. Duff, a local farmer on South Bass Ialand, told me 171 of his experience with grackles and their affinity for grapes. One day, a few years ago, when automatic scar­ ing devices were not available and protection depended upon the patrol with a shotgun, he was called off the island on urgent business for half a day during the grape ripening season and returned to find his arbors dark with grackles. The destruction they caused was not min­ imal for it was estimated that one-half of the total crop was destroyed in that one afternoon the grapes were left unprotected.

Nestling food types The greatest difference between the food types eaten by the adults and that fed to the nestlings was found to be a reversal in the percentages of animal versus plant food consumed. As was noted earlier, plant mater­ ials were found to be the primary food item for adults. The nestlings received animal foods almost exclusively, especially during the early days of nest life. No nest­ ling stomachs were examined, the food identification was made through a telescope at the nest site. This proce­ dure made quantitative studies Impossible, but Beal (1900) investigated the stomachs of 456 nestlings and found 7^.^ per cent animal matter and 25.6 per cent plant. 172 The Put-in-Bay nestlings from Nests 8-64 and 20-64 received the following food items in this approximate order of frequency: Ephemeroptera, Tricheoptera, Diptera, Coleoptera larvae, Lepidoptera adults and larvae, Araneida, Oligochaeta, P u m u s sp., fish carrion, bread and a small snake. In addition to these food items Beal (1900) mentions: Orthoptera, Hymenoptera, Hemiptera and com. It is probable that these items missed detection and would have been found in the Put-in-Bay birds' stomachs had they been examined. Spiders were first identified on the first day of nest life, and were delivered to the young well crushed during the early hours of nestling life. On the second day of nest life, mayflies were stuffed into the nest­ lings' open mouths and this continued to be the major food item throughout nest life. Late spring and early summer are times of good mayfly hatches and the mayfly supply must be an important factor in establishing them as the primary food item. The same conditions sore present to enable the caddisfly to be the second most important food item. Later in June while Nest 20-64 was under ob­ servation the mayfly supply diminished and adult grackles were noticed taking flesh from dead perch (Perea flavesoens) along the Lake Erie shore. The adults were followed to 173 the nest and seen feeding this carrion to the nestlings. This food type determination through the telescope would have been impossible had I not seen the source of the food supply. Hamilton (1951) examined 130 Bronzed Grackle nest­ ling stomachs and feces at Ithaca, New York over a three year period. He found the animal remains to make up 89.I per cent of the total volume, the plant remains 6.4 per cent. Table 32 gives a summary of the per cent occurrence in the stomachs and a percentage by volume.

Unusual feeding behavior The Common Grackle has been seen as a vertebrate predator and carrion eater in addition to its dominant omnivorous feeding habits discussed above. The unusual feeding behavior most often reported in the literature is with respect to fish. Cottam (1943) reported grackles feeding on dead fish parts at the base of a power dam with the dexterity of typical water birds. Reports of the hovering, gull-like fishigg behavior were published by Townsend (1919) who found the grackles eating live stickleback (Gasterosteus aculeatus) from the Charles River in Boston, Cahalane (1944) reported grackles taking small dead perch (Perea flavesoens) from the sur­ face of Lake Michigan. I observed grackles picking at 174-

TABLE 32

Food of 130 nestling Bronzed Grackles in the Ithaca, New York, region, May and June, 1947-1949. Hamilton (1951)

Food Percent occurrence Percent by volume

Insects 84.6 48.2 Earthworms 18.5 10.9 Amphibians 10.8 8.3 Fish 70.8 6.6 Spiders 24.6 5.4 Green grass 9.2 3.2 Millipedes 6.1 0.8 Grain 4.6 1.4 Sowbugs 4.6 3.4 Fruit 4.6 1.8 Mammals 1.5 0.3 Molluscs 2.3 5.2 Grit 17.0 4.3 175 dead, perch on Lake Erie’s shore at Put-in-Bay, Ohio and subsequently fly with it to their nestlings. Also the eyes were removed from dead perch and eaten by the adult during my observations. Follett (1957) observed grackles taking live emerald shiners (Notropls atherlnoldes atherlnoldes) from the Niagara River below the Falls. Hamilton (1951) during his nestling food study discussed earlier, noted the grackles picking up small dead alewives (Alosa pseudoharengus) from the shore of Cayuga Lake at Ithaca, New York, and feeding them to the nestlings.

Beeton and Wells (1957) saw a female Bronzed Grackle take live emerald shiner (Notropls atherlnoldes acutus) and feed them to her young. They give this description of the capture: "The bird flew back and forth eight to ten feet above the water, then upon sighting a minnow it dipped down, hovered immediately above the fish and captured it with a quick thrust of the beak.” Although I have never seen a grackle attack and kill other birds, their eggs or frogs, there are a few literature reports. Attacks of male Purple Grackles on young Pheasants in rearing cages is discussed by Foster (1927). The graokles entered the enclosure and beheaded the pheasants, killing three hundred before they were stopped. Davis (19*J4) saw a Purple Grackle stalk a House Sparrow (Passer domestlous). The sparrow was seized by Its beak and dashed to the pavement, Its viscera eaten by the grackle. Similar behavior was observed by Christofferson (1927) when a Bronzed Grackle pecked a Pine Siskin and B a m Swallow In the head, leaving only part of the skull Intact and the brains gone. Ernst (19*l4) noted the Bronzed Grackle near Olean New York, attack small leopard frogs (Rana plplens) in a pond by piercing their soft throat or eyes with the bill They would fly with their prey to a tree and grasp the frog with one foot and tear its flesh with their bill. Parasitism

This section is concerned with the internal and external parasites of the Common Grackle (Qulsoalus qulsoula versicolor)• The data collected at Put-in-Bay, Ohio are supplemented by the literature. The methods used are discussed in the materials and methods section. Parasitism of nestlings by calliphorid flies has been reported in the literature, but I can find no reference to bird egg parasitism by maggots of the Family Calliphorldae. Neff (19^5) studied nestling Infestations by maggots of the genus Protocalllphora in the Mourning Dove (Zenaldura maoroura), House Sparrow (Passer domestlous). Mockingbird (Mlmum polyglottos). Shrike (Lanius ludovlolanus), and Western Kingbird (Tyrannus vertioalis). The maggots infested the nestlings* ears, nostrils, wings, tall, legs and were found among the pinfeathers. Blair (1931) found Phaenloia serloata maggots in the nests of the Hook (Corvus frugllegus) and Johnson (1932) found the same species in the nests of the B a m Swallow (Hirundo ervthrogaster). A compilation of bird nest inseOt

parasites is found in Hicks (1959)*

177 1?8 During the measurement of non-fertile grackle eggs, I accidently dropped an egg and found maggots swarming in the yolky albumen. These maggots were found in rancid eggs in which no cracks were noted with the unaided eye. The grackle eggs were placed in a wire cage in a dark, humid room. Several adults emerged which were collected and pinned for identification. All flies identified were one species - Phaenicla serioata Meigen, Family Calll- phoridae. The determinations were made by Mr. C.W. Sabrosky of the United States Department of Agricultural Research Servioe, Beltsville, Maryland, Examination of 15 intestines made under the dissection microscope found 13 Infested with one species of spiny- headed worm. The maximum number of worms in each intestine was four and the mean number was two per intestine for the 13 infested guts. No flukes, tapeworms or nematodes were observed. Findings of others are included in the summary at the end of this section. Fifteen specimens were washed for ectoparasites, which were collected, mounted and identified by experts credited in the summary. Ten species of Aoari are recorded. The nomenclature used in the summary which follows is based upon the latest literature found. In the case of the Acari, recent nomenclatorial revisions were made at The Institute of Acarology in Wooster, Ohio by Mr. Donald E. Johnston, Curator. 179

Acanthocephala

Medi orhynchus grandis Van Cleave, 1916 Site of Infection - Intestine Classification according to Van Cleave (19*^8) and Chandler (1961). Order - Archiacanthooephala Meyer, 1931* Family - Gigantorhynohus Hamann, 1892. Genus - Mediorhynchus Van Cleave, 1916. Authority - G.W. Welker (1962) and G.R. Spory (1963).

Prosthorhvnchus formosus (Van Cleave, 1918) Travassos, 1926. Site of Infection - Intestine

Classification according to Yamagutl (1963). Order - Echinorhynchidea Southwell et Macfie, 1925. Family - Plagiorhynchidae Golvan, i960. Genus - Plaglorhynohus Luhe, 1911. Authority - Present study. Det. by W.W. Becklund. 180

Cestoda

Orthoskr.1 ablnla qulscall (Hodgers, 1941) Spassky, 1947. Site of Infeotion - Small intestine. Classification according to Yamaguti (1959)* Order - Cyclophyllidea Braun, 1900. Family - Dllepididae Railliet and Henry, 1909• Genus - Orthoskr.1 ablnla Spassky, 19^7• Authority - G.W. Welker (1962).

Parloterotaenia parlna (Dujardin, 1945) Fuhrmann, 1932. Site of Infection - Small intestine. Classification according to Yamagutl (1959). Order - Cyclophyllidea Braun, 1900. Family - Dllepididae Railliet and Henry, 1909• Genus - Parloterotaenla Fuhrmann, 1932. Authority - G.W. Welker (1962). Trematoda

Consulcuum loterldorum Denton and Byrd, 1951. Site of Infection - Gall bladder. Classification aooording to Yamagutl (1958) Order - Dlgenea Van Beneden, 1858, Family - Dicroooellidae Odhner, 1911. Subfamily - Dioroooelllnae Looss, 1899 Genus - Consulcuum Bhalerao, 1936. Authority - G.W. Welker (1962).

Consulouum macrorohIs Denton and Byrd, 1951* Site of Infection - Gall bladder. Classification according to Yamagutl (1958) Order - Dlgenea Van Beneden, I858. Family - Dicrocoelildae Odhner, 1911. Genus - Consulouum Bhalerao, 1936. Authority - G.W. Welker (1962).

Brachyleo1thum amerlcanum Denton, 19^5. Site of Infection - Small intestine. Classification according to Dawes (19^6). Order - Dlgenea Van Beneden, 1858. Family - Echinostomatldae Looss, 1902. Genus - Echlnostoma Rudolphi, 1809. Authority - G.W. Welker. (1962). Eohlnostoma revolutum (Froelich, 1802) looss, 1899* Site of Infection - Small intestine. Classification according to Dawes (19^) • Order - Dlgenea Van Beneden, 1858. Family - Echinostomatidae Looss, 1902. Genus - Echlnostoma Rudolphi, 1809* Authority - G.W. Welker (1962).

Crepldostomum oooperl Hopkins, 1931* Site of Infection - Gall bladder. Classification according to Yamagutl (1958). Order - Dlgenea Van Beneden, 1858. Family - Allocreadildae Stosslch, 1903* Subfamily - Crepldostomum Dollfus, 1951. Genus - Crepldostomum Braun, 1900.

Authority - G.W. Welker (1962). 183 Nematod®

D1spharynx spiralis (Molin, 1858) Skrjabin, 1916. Site of Infection - Mucosa of digestive tract. Classification according to Cram (192?). Suborder - Spirurata Railliet and Henry, 1915. Family - Acuariidae Seurat, 1913* Subfamily - Acuarilnae Railliet, Henry and Sisoff, 1912. Genus - Pispharynx Railliet, Henry and Sisoff,1912.

Authority - G.W. Welker (1962).

Caplllarla qulscall Read, 19*1-9. Site of Infection - Lower intestine. Classification according to Chitwood and Chitwood (1950). Superfamily - Trichuroidea Railliet, 1916. Family - Trichuridae Railliet, 1916. Genus - Caplllarla Zeder, 1800. Authority - G.W. Welker (1962).

Splendldofliarla quisoall Odetoyinbo and Ulmer, i960. Site of Infection - Pia mater of cerebral hemisphere. Classification according to Anderson and Chabaud, 1959* Family - Onchocercldae Anderson and Chabaud, 1959* Subfamily - Splendldofilaria Chabaud and Choquet,1953* Genus - Splendldofilaria Skrjabin, 1923* Authority - G.W. Welker (1962). 184

Acari

Paraneonyssus loterldlus Strandtmann and Furman, 1956. Site of Infeotion - Nasal passages. Classification according to Strandtmann and Wharton (1958). Order - Acari Suborder - Mesostigmata Canestrini, 1891. Family - Rhinonyssidae Trouessart, I895. Genus - Paraneonyssus Castro, 1948. Authority - Present study. Det. by R.W. Strandtmann.

Omlthonyssus sylviarum (Canestrini and Fanzago, 1877). Site of Infection - Skin and feathers. Classification according to Strandtmann and Wharton (1958). Order - Acari Suborder - Mesostigmata Canestrini, 1891. Family - Laelaptidae Berlese, I892. Genus - Ornlthonyssus Sambon, 1928. Authority - Present study. Det. R.W. Strandtmann. 185

Androlaelaps oasalls (Berlese, 188?) Bregetova, 1956. Site of Infection - Feathers. Classification according to Strandtmann and Wharton

1958). Order - Acari Suborder - Mesostigmata Canestrini, 1891. Family - Laelaptldae Berlese, I892. Genus - Androlaelaps Berlese, 1903. Authority - Present study. Det. by E.W. Baker.

Heamaphysails leporlspolustrls (Packard, I869). Site of Infection - Skin. Classification according to Nuttall and Warburton

(1911). Order - Acari Suborder - Ixodldes Leach, 1815.

Family - Ixodidae Murray, 1877* Genus - Haemaphysal1s Kock, 184^. Authority - Present study. Det. by C.M. Clifford. 186

Mesalges sp. Site of Infection - Feathers. Classification according to Zumpt (1961). Order - Acari Suborder - Aoaridei Latreille, 1802. Family - Analygesldae Megnin, 1800. Genus - Mesalges Trouessart, 1888. Authority - Present study. Det. by W.T. Atyea.

Proctophyllodes egglestonl Spory, 19^5 • Site of infection - Skin and feathers. Classification according to Zumpt (1961). Order - Acari Suborder - Acaridei Latreille, 1802. Family - Proctophyllodidae Megnin and Trouessart, I883. Genus - Prootophyllodes Robin, 1868. Authority - Present study. Det. by W.T. Atyeo. 187

Harpyrhynohus sp. Site of Infection - Skin and feathers. Classification according to Baker and Wharton (1952). Order - Acari

Suborder - Trombidlformes Reuter, I9 0 9 . Family - Harpyrhynchidae Dubinin, 1957* Genus - Harpyrhynohus Megnin, 1877. Authority - Present study. Det. by G.M. Clark.

Syrlngophllus elongatur Ewing, 1911 - (icteridae Clark, Site of Infection - Feather quills. 196*0. Classification according to Baker and Wharton (1952). Order - Acari

Suborder - Trombldifomes Reuter, I9 0 9 . Family - Syrlngophilidae Lavoipierre, 1953. Genus - Syrlngophllus Heller, 1880. Authority - Present study. Det. by G.M. Clark. 188

Eutrombloula alfreddugesl (Oudemans) Site of Infection - Skin. Classification according to Baker and Wharton (1952). Order - Acari Suborder - Trombidlformes Reuter, 1909. Family - Tromblculidae Ewing, 1944.

Genus - Eutrombicula Ewing, 1938. Authority- - Present study. Det. by J.M. Brennan. Summary

1. This study was completed In 1964 and 1965 at the Franz Theodore Stone Laboratory of The Ohio State Univer­ sity, Put-in-Bay, Ottawa County, Ohio. 2. There are three subspecies of the Common Grackle (Quiscalus qulsoula) t Florida Grackle (£.3,. qulsoula); Purple Grackle (Q.q. stonel); and Bronzed Grackle (Q.cj.. versicolor). The latter subspecies is the subject of this investigation, although called the Common Grackle throughout the dissertation. 3. The scientific name of the Common Grackle subspe- cies Qulscalus qulsoula versicolor Vieillot has remained stable since 1948. 4. There is significant individual variation in the length of wing and tail feathers among specimens of the same sex. 5. Sexual dimorphism is pronounced in body size and color differences with male wing feathers 11 per cent longer and tail feathers 13 per cent longer than the fe­ male^. Feather coloration in the female is duller with the metallic hues more subdued. 6. The Common Grackle has one complete moult per year and the definitive plumages are acquired after the bird's

189 190 first year. Eye color becomes lighter as the feather color becomes darker during the first year. 7. All subspecies of the Common Grackle are dis­ tributed through the east and central United States, to the east and central portions of south Canada. 8. The unusual voice of the grackle played an im­ portant role in courtship, species recognition, communi­ cation and individual recognition among pairs. 9. Voice development is slow, not reaching full maturity until the second year. The nestlings* first sounds were noted on the second day of nest life and in­ creased in intensity until the eleventh day when they were capable of giving the "intense alarm". 10. Pair formation begins shortly after the arrival of the females to the breeding grounds and continues, in some cases, into the nest building phase. The transition from the winter flocking behavior to pair formation is quite gradual. 11. The pair bond is achieved through two stages: (1) maintenance of contact between Individuals of both sexes and (2) acceptance of the male by the female. 12. Multi-male flights are a conspicuous part of pair formation behavior. Plights Involving as many as five males and one female are common earlier in the season, with two males in flight with a female more common toward the end of pair formation. The female Initiates most 191 flights and the male will strive to achieve a dominant perch at the end of the flight. 13. Keeling of the tail is a display used by the male during pair formation and paired flights. It Is strongest in the early morning and early in the breeding season. The intensity decreases after incubation has begun. 14. Copulatory behavior was noted at least once in the early morning and once in the evening near the nest site during the days that eggs were laid. 15. All nests in the Put-in-Bay, Ohio colony were located in Juniper (Juniperus virgin!ana) with the excep­ tion of one which was found in boxwood, an ornamental. The median nest height was 20 feet. 16. Length of time for nest building varied from seven consecutive days to a non-conseoutive eight weeks. A count of 124 visits to one nest during the nest building period is an estimate of the total visits involved in the con­ struction of one nest. 17. The first egg was laid three days after the nest was completed in three closely watched nests. 18. There is a limited territory around the nest, the defense of which is strongest during the early stages of the breeding cycle. The object of the male's defense switches from the nest site to the female and young dur­ ing the progressive stages of the breeding cycle. 192 19. Measurements of 256 eggs were obtained from three sources which gave a mean length of 28.14 millimeters, a mean breadth of 20.27 millimeters and a mean elongation of 1.38 millimeters. 20. Egg laying times at 13 nests were determined. The eggs are laid at approximately 24 hour Intervals and between the hours of 0620 and 1000. 21. Mean clutch size of 4.7 was determined from 187 nests and 881 eggs whioh included the Put-ln^Bay, Ohio nests supplemented by literature reports. 22. Incubation is the responsibility of the female, the male's role restricted to guarding the nest and maintain­ ing the pair bond by his presence near the nest site. 23. Incubation constancy or the percentage of ob­ served time the female spends on the nest was calculated at eight nests. Prom a total observed time of 529 hours and 31 minutes, the constancy for all nests was 65 per cent. If only those nests which hatch eggs are considered, the constancy was 73 per cent. This indicates that fe­ males are more attentive to successful nests than to un­ successful ones. 24. Attentive behavior of Common Grackle females dur­ ing the normal day one to day 13 incubation period is evenly distributed during morning, mid-day and afternoon hours. 193 25. There was a negative correlation between air temperature and per cent constancy. The higher the air temperature, the lower the constancy and the lower the air temperature, the higher the constancy. 26. The length of the incubation period was deter­ mined to the nearest hour from seven nests. The mean length was 13 days and 4 hours, the shortest time being 12 days and 8.5 hours and the longest time 14 days and 2.5 hours. 27. Brooding is the responsibility of the female, al­ though a male did brood at one nest for 10 minutes each day on the third, fourth and fifth days of nest life. 28. There was considerable variation in the amount of time spent brooding by the female at two nests. In one case brooding stopped on the eighth day of nest life, but in another nest continued until fledging. 29. Both the male and female participate in feeding activity. The combined feeding activity by both male and female adults at two nests was comparable. A 2.55 feedings per nestling per hour was calculated for one nest and 2.10 feedings per nestling per hour for another. 30. Feeding activity of the male and female is vari­ able from nest to nest. In one nest, the male was slightly more active in feeding than the female, but in another nest the male was almost totally inactive, the female accomplishing most of the feeding tasks. 194

31. The male and female shared in fecal sac removal, but the female was responsible for parasite removal and nest repair. Pecal sacs were emitted by nestlings at more than half the visits by the adults. 32. Pecal sacs were eaten in a greater percentage during the early days of nest life, with a decrease in frequency later. The female at one nest disposed of 45 per cent of the fecal sacs and at another nest 82 per cent. 33. The differences between activity of the sexes at two nests are stable regardless of the type activity. The relative inactivity of the male at Nest 20-64 is reflected in feeding, brooding and nest maintenance behavior. 34. Total dependency of the young grackle on the adult for feeding appears to last two weeks past fledging. 35. At least one nestling fledged from ten nests of 24 active nests for a nest success of 41 per cent. A nest­ ing success of 47 per cent is obtained when this study, Eyer (195^) and Petersen and Young (1950) studies are com­ bined. 36. Nest failures at the Put-in-Bay, Ohio colony in the order of frequency of occurrence are due to: human interference, weather, infertile eggs and predators. 37. The mean hatch weight for five nestlings was 6.8 grams. The mean weight on the last day of nest life for three nestlings was 73*3 grams. Daily weight gains averaged 5.04 grams over a 12 day period. 195 38. The tarsus and wing of four nestlings grew very nearly the same rate during the 12 day nestling period, 2.45 millimeters per day and 2.43 millimeters per day, respectively. The bill grew at the slowest rate of 0.68 millimeters per day and the ninth primary the most rapidly at 3*00 millimeters per day. 39• There were 3315 feathers counted in all tracts on a 12 day old nestling. The ventral tract had the most feathers with 1034, the caudal tract the least with 94 feathers. 40. The first migratory wave arrives in central Ohio from the wintering grounds in February and departs in November. A check of banding records indicates that a northward post-breeding dispersal is probable for a small proportion of the population. 41. The largest wintering populations of Common Grack- les were found in Norfolk County, Virginia in i960 and I96I,

Rome, Georgia in 1962 and Little Rock, Arkansas in 19^3* 42. Recoveries of Ohio banded grackles indicate that the greatest number winter in Tennessee and Alabama, with some recorded from Mississippi, Louisiana, Kentucky, Indi­ ana, Georgia, Florida, Arkansas and Texas. 43. The longevity record for the Common Grackle is 17 years (Bent, 1958:404). The oldest Ohio banded grackle found in banding files of the U.S. Fish and Wildlife

Service was 10 years. 196

44. The grackle is omnivorous and eats what is avail­ able at the time. Examination of 24 stomachs revealed that the male's stomach contents hsvd a dry weight for plants of 0*3576 grams, the female's 0.2924 grams and the immatures 0.2830 grams. 45. Plants accounted for 93*2 per cent of the total dry food weight, animal remains were 4.3 per cent and stones plus other miscellaneous items 2.5 per cent. 46. The grackle in company with Red-wings is a ser­ ious threat to the grape crops on South Bass Island, with carbide scaring devices and personal patrols the farmer's defense. 47. The nestling receives more animal matter than plant. At Put-in-Bay, Ohio, the major food item was the mayfly, with the caddlsfly and midges the next most abun­ dant food. 48. The grackles obtained flesh from dead perch for nestlings' food and were seen eating the eyes from re­ cently dead yellow perch. 49. Maggots of the species Phaenlcla- serloata. Family Calliphoridae were found in rancid grackle eggs obtained from abandoned nests. 50. An investigation of ecto-and endoparasites yielded one species of spiny-headed worm from the gut and ten species of Acari from feathers and skin. Appendix

197 198

TABLE 33

Daily morning and afternoon temperatures at Put-in-Bay, Ohio.

Day, Temp. °F Day, Temp. °F Month, 0700 hrs. 1 6 0 0 hrs. Month, 0 7 0 0 hrs. 160 0 hrs, Year 1961* 1965 1961* 1965 Year 1961* 1965 I96I* I9 6 !

1 May 51 1*9 51 53 1 June 57 71* 61 71 2 May 52 60 53 55 2 June 6 0 61* 62 55 3 May 55 65 52 76 3 June 55 68 63 57 1* May 58 52 57 50 1* June 69 59 60 59 5 May 61 1*8 71 51 5 June 6 0 61 6 May 68 53 78 56 6 June 6 0 65 7 May 60 76 77 61 7 June 63 70 8 May 61* 73 71 82 8 June 6U 68 9 May 67 70 70 80 9 June 78 86 10 May 6h 65 65 71* 10 June 80 65 11 May 67 63 66 65 11 June 65 63 12 May 59 60 57 58 12 June 62 69 13 May 59 61 57 52 13 June 79 80 11 May 1*9 62 58 60 Hi June 69 70 15 May 63 65 71 83 15 June 79 71 16 May 67 70 70 77 16 June 55 63 17 May 62 58 69 60 17 June 65 67 18 May 62 57 72 61 18 June 72 81 19 May 75 65 71 61* 19 June 83 88 20 May 50 57 58 60 20 June 76 77 21 May 59 6 0 63 67 21 June 78 70 22 May 70 68 81* 69 22 June 72 72 23 May 75 56 81 58 23 June 77 85 21* May 75 57 68 60 2h June 75 68 25 May 62 65 71 80 25 June 71* 73 26 May 6k 75 66 70 26 June 80 89 27 May 63 68 65 70 28 May 58 60 61 59 29 May 50 50 58 55 30 May 61 62 6 0 59 31 May 58 68 58 65 199 TABLE 3k

Measurements of Quiscalus quiscula versicolor eggs from the Jr. B.R. Bales Collection in The Ohio State Museum, the Preston Laboratories and from Put-ir-Bay, O'- io. (in millimeters)

Egg source Dr. B.R. Bales Collection ; Ohio Egg measurer Dr. B.R. Bales.

No. of Eggs Length (mm) Breadth

1 # 27.25 19-25 2 28.25 19.00 3 27.00 19.25 k 27.50 19.00 5 * 26.75 10.50 6 27.00 18.75 7 27.75 19.50 p> 26.50 18.75 9 # 27.75 20.00 10 27.50 20.00 11 26.00 20.50 12 28.00 10.75 13 * 20.25 20.25 lk 20.75 20.00 15 29.00 20.25 16 20.25 20.00 17 * 23.75 19.75 18 27.75 19.75 IQ 29.00 19.75 20 28.25 10.75 21 # 2k. 75 20.00 22 2k. 50 19.75 23 2k. 50 19.25 2k 2k. 25 10.50 25 # 29.50 20.50 26 29.75 20.75 27 28.50 19.75 28 28.25 20.25 2P 27.00 20.50 3 0 # 28.50 20.50 31 28.50 19.75 32 28.00 20.50 33 28.50 19.75 3k 28.50 19.75 35 # 29.75 19.50 36 29.50 10.00 37 30.50 19.50 38 30.75 19.25 30# 25.75 19.00 ko 26,25 18.75

# = Beginning of clutch TABLE 3k (condt.) 200

: o. of Eggs Length (mm) Breadth (mm)

111 27.00 10.25 112 2k.SO 19.50 113 * 25.75 20.00 hh 25.50 20.00 kS 26.00 19.50 k6 27.25 19.75 hi 25.25 19.50 i.8 * 26.25 19.50 U9 27.00 19.50 50 27.50 19.00 51 27.25 19.00 52 27.75 18.50 53 * 26.25 19.50 511 27.25 19.75 55 26.25 19.75 56 26.25 19.50 57 * 23.75 19.25 58 28.75 19.75 5° 27.50 13.75 60 32.25 18.25 61 * 27.25 21.25 62 28.50 21.25 63 27.75 21.25 6h 27.50 20.75 65 27.00 20.75 66 20.25 21.50 67 29.50 22.00 68 2o.2 5 21.75 69 28.25 22.00 70 29.50 21.75 71 * 28.00 20.75 72 28.00 20.50 73 27.25 20.00 Ik 28.50 21.25 IS 29.50 21.25 76 29.25 20.25 77 29.25 21.00 78 29.25 21.00 79 29.25 20.50 80 * 26.75 21.50 81 27.00 21.00 82 27.25 21.00 83 26.50 20.75 8ii 27.25 20.75 85 * 20.25 20.50 86 28.00 21.00 87 29.25 20.25 88 28.00 20.75 89 23.50 20.25 o o 41- 30.25 20.50 TABLE 3b (condt.) 201 ho. of Eggs Length (mm) Breadth (mm)

01 23.25 10.50 02 29.00 20.50 °3 30.75 10.75 °b :: 29.50 21.00 05 30.50 20.25 «6 27.75 20.75 97 29.75 21.75 98 28.75 20.75 99 20.50 20.75 100 23.75 20.50 101 28.25 20.25 102 * 28.00 21.00 103 26.25 . 20.00 10b 27.50 21.00 105 27.75 20.50 106 28.25 20.75 107 * 25.50 19.50 108 27.00 18.75 109 28.00 10.50 110 27.75 19.25 111 27.25 18.75 112 27.00 20.50 113 26.75 20.50 11U 2 7.00 20.50 115 25.00 19. 75 116 25.75 20.00 117 28.50 20.00 118 27.50 20.00 n o 28.00 20.75 120 27.50 21.00 121 28.25 19.50 122 28.50 20.75 123 28.50 20.50 12b 28.25 20.25 125 * 27.26 20.50 126 27.25 20.50 127 28.00 20.00 128 27.00 19.75 129 28.25 20.00 130 29.50 20.25 131 27.75 20.00 132 27.00 20.00 133 - 20.25 20.50 13b 27.25 20.00 .135 29.50 21.00 136 28.25 21.50 TABLE 3h (condt.) 202

Mo. of Kftgs Length (mm) Bre-pdth (mm)

137 28.29 21.50 133 28.73 21.25 13? * 23.73 20.50 1110 27.73 20.50 1111 27.00 20. 75 m 2 27.00 20.50 1)4.3 *- 27.73 20.50 lhh 28.30 20.00 ihs 27.50 20.50 i36 * 28.00 20.25 U 7 27.00 20.00 1)48 27.30 19.50 Ih0 23.75 19.50 mo 26.50 20.25 m i *- 27.00 Id. 75 152 25.75 19.75 153 27.00 19.25 mu 26.75 19.75 155 * 27.75 20.75 m 6 27.75 20.25 157 28.25 20.00 l?8 27.70 20.00 l$o 28.25 20.75 160 44 27.25 19.50 161 26.75 Id. 00 162 28.50 17.50 163 27.75 20.00 16U 27.25 19.75 163 28.25 20.50 166 28.50 20.75 167 28.75 20.50 168 28.50 20.50 160 29.25 19.50 170 28.75 20.75 171 30.75 20.75 172 29.50 20.50 173 28.00 19.50 17U 29.50 20.25 173 * 27.25 19.75 176 27.50 19.25 177 27.50 20.25 178 27.75 19.25 179 27.75 20.25 180 26.75 20.25 181 28.50 19.50 182 28.50 20.25 183 2d.50 20.00 18U 28.50 20.25 183 28.75 70.50 TABLE 3U (condt.) 203

No. of Eggs Length (mm) Breadth 186 29.50 19.50 187 29.25 19.50 188 29.75 19.50 189 29.00 19.00 190 28.50 19.50 191 27.75 20.75 192 28.50 20.25 193 27.25 20.75 19U 27.25 20.50 195 26.00 20.75

Egg source Dr. B.R. Bales Collection 3 Pennsylvania Egg measurer Dr. B.R. Bales.

No. of Eggs Length (mm) Breadth

1 * 28.00 20.50 2 28.75 20.75 3 28.25 20.25 h 28.75 19.75 5 29.25 20.00 6 -;<• 28.75 20.00 7 29.50 21.00 8 28.75 20.75 9 29.25 21.25 10 28.75 20.75 11 -if- 29.75 20.75 12 29.75 20.50 3.3 29.50 20.50 1U 31.00 20.75 15 30.00 20.75 16 # 26.00 19.75 17 26.25 20.50 18 26.75 20.25 19 27.00 20.00 20 27.00 20.75 21 -if- 29.00 20.50 22 29.50 20.50 23 27.00 20.00 2h 27.75 20.50 25 27.25 20.25 204 TABLE 31* (condt,)

Egg source Dr. B.R. Bales Collection ; Minnesota Egg measurer Dr. B.R. Bales. lo, of Eggs Length (mm) Breadth (mm)

1 * 27.75 18.75 2 28.25 19.25 3 28.75 18.50 1* 28.75 . 19.25

Egg source Preston Laboratories 3 Carnegie Museum 3 Pennsylvania Egg measurer Frank W. Preston Wo. of Eggs Length (mm) Breadth (mm) (1 egg per clutch)

1 28.00 20.69 2 30.88 22.00 3 27.53 21.08 1* 29.31* 22.15 5 30.38 21.13 6 27.05 21.61 7 31.12 21.76 8 30.22 20.12 9 30.13 20.30 10 28.51* 21.15

Egg source Preston Laboratories 3 Carnegie Museum 3 Ohio Egg measurer Frank W. Preston

No. of Eggs Length (mm) Breadth (mm) (1 egg per clutch)

1 27.85 21.08

Egg source Pr ;ston Laboratories 3 Brandt Collection 3 Ohio Egg measurer Mo. of Eggs Length (mm) Breadth (mm) (1 egg per clutch)

1 30.73 21.68 2 23.30 20.07 J> 27.92 21.72 1* 29.81* 21.12 205 TABLE 3U (condt.)

Sfifr source Preston Laboratories ; Brandt Collection1 Magdalen Is.(Que.) Eg;, measurer Frank W. Preston

No. of Eggs Length (mm) Breadth (mi,-.) ''I egg per clu tch )

1 30.SU 20.61 2 28.8U 21.99 3 29.36 21.37 h 20.38 21.06

Egg source Preston Laboratories; Brandt Collection ; Alberta Egg measurer Frank W. Pres t'in

Fo. of Eggs Length (mm) Breadth (mm) (1 egg per clutch)

1 30.35 20.61 2 . 29.70 20.70

Egg source Put-in-Bay, Ohio Egg measurer G.R. Maxwell, I I

No. o f Eggs Length (mm) Breadth (mm)

1 2U.5Q 19.Uo 2 30.80 21.80 3 29.70 21.80 k 30.60 20.80 5 30.90 21.20 6 30.60 21.80 7 2Q.90 21.60 8 27.80 21.70 9 27.00 21.SO 10 30.So 21.Uo 11 32.00 21.60 TABLE 35

Summary of feedings per nestling per hour by male and female adult Common Grackle (Quiscalus quiscula versicolor) from Nest 8 — 61j. (5 nestlings) and Nest 20-61; (3 nestlings) at Put-in-Bay, Ohio.

Days of Nest Life

Nest 1 2 3 U 5 6 7 8 9 10 11 12

8-6U

Total feedings/nestling/hr. 9 0.28 0.21; 0.1*1; 0.60 0.67 0.72 0.59 1.12 O .96 1.81 1.32 1.08 Total feedings/nestling/hr. c1 0.00 0.73 0.69 0.83 0.69 0.90 0.76 1.1*1 l.lil* 1.36 1.18 1.08 Total feedings/nestling/hr. 9 d 0.28 0.97 1.13 1.1*3 1.36 1.62 1.35 2.53 2.1*0 3.17 2.50 2.16

20-61;

Total feedings/nestling/hr. 9 1.13 0.62 1.06 1.31 1.2? 1.32 1.31 1.21 1.96 1.35 1.33 2.21; Total feedings/nestling/hr. d 0.00 0.00 0.09 0.05 0.30 0.1;9 0.53 0.12 0.35 0.73 l.oU 0.16 Total feedings/nestling/hr. 9 d 1.13 0.62 1.15 1.36 1.57 1.81 1.81; 1.33 2.31 2.08 2.37 2«li0

t\> o os TABLE 36

Disposition of time the female adult Common Grackle (*Quiscalus quiscula versicolor) spent at Nests 8—6/4. and 20-6k, Put-in-Bay, Ohio;during the daylight hours.

Days of Nest Life

1 2 3 h 5 6 7 8 9 10 11 12

Nest 8-6ii

Percent of time brooding hS 5i la 31 17 7 5 0 0 0 0 0 Percent of time away from nest 52 hi 55 6k 77 81 82 90 91 82 89 92 Percent of time feeding young 3 2 h 5 6 6 5 8 7 13 9 8 Percent of time caring for nest 0 0 0 0 0 6 8 2 2 5 2 0

Nest 20—61i

Percent of time brooding 38 58 38 38 22 55 31 ho 31 26 8 28 Percent of time away from nest 60 ill 59 61 73 lil 62 57 58 71 75 61 Percent of time feeding young 2 1 3 1 3 3 3 3 h 3 3 6 Percent of time caring for nest 0 0 0 0 2 1 h 0 7 0 Hi 5

ro o TABLE 37

Brooding and feeding of Common Grackle (Quiscalus quiscula versicolor) nestlings from Nest 8-61*, Put-in-Bay, Ohio.

Days of Nest Life

1 2 3 h 5 6 7 8 9 10 11 12

Total time observed (hr & min) 3 Ok 5:1*1* l*:0l* 8:39 7:30 5:35 5:1*6 7:37 5:12 2:1*6 5:06 2:19 Total visits/hr. 9 1*.20 3.66 3.93 1*.39 3.U7 1*.1*8 3.99 h.99 1*.1*2 7.9l* 5.1*9 1*.31 Feeding only visits/hr. 9 0.00 0.00 0.25 0.92 1.87 2.87 2.60 1* .33 3.85 7.22 5.29 1*.31 Brooding only visits/hr. 9 2.80 2.27 1.72 1.39 0.13 0.00 0.17 0.13 0.19 0.00 0.00 0.00 Feed and brood visits/hr 9 1.1*0 1.22 1.97 2.08 1.1*7 0.72 0.35 0.13 0.00 0.00 0.00 0.00 Total feeding visits/hr. 9 l.ho 1.22 2.21 3.01 3.33 3.58 2.95 1*.1*6 3.85 7.22 5.29 li.31 Total visits/hr. d 0.00 3.81* 3.69 1*.28 3.60 1*.66 3.81 5.77 5.77 5.1*2 1*.71 li.31 Feeding only visits/hr. d 0.00 3.66 2.95 I*.l6 3.33 1*.1*8 3.81 5.61* 5.77 5.1*2 1*. 71 li.31 Brooding only visits/hr. d 0.00 0.00 0.25 0.12 0.13 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Feed and brood visits/hr d 0.00 0.00 0.1*9 0.00 0.13 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Total feeding visits/hr. d 0.00 3.66 3.1*1* U. 16 3.1*7 li.liQ 3.81 5.61* 5.77 5.1*2 1*. 71 1* .31 Total visits/hr. 9 d ' l.ho 7.50 7.62 8.67 7.07 9.11* 7.80 10.76 10.19 13.36 10.20 8.62 Feeding only visits/hr. 9 tf 0.00 3.66 3.19 U.86 5.20 7.35 6,1*1 9.97 9.62 12.61* 10.00 8.62 Brooding only visits/hr. 9 d 2.80 2.27 1.97 1.50 0.27 0.00 0.17 0.13 0.19 0.00 0.00 0.00 Feed and brood visits/hr . 9 d l.ho 1.22 2.1*6 2.08 1.60 0.72 0.35 0.13 O.C'j 0.00 0.00 0.00 Total feeding visits/hr. 9 d 1,1*0 1*.8? 5.65 7.17 6.80 8.06 6.76 10.10 °.62 12.61* 10.00 8.62 208 TABL55 38

Brooding and feeding of Common Grackle (Quiscalus quiscula versicolor) nestlings from Nest 20-6h} Put-in-Bay^ Ohio.

Days (:>f Nest Life

1 2 3 h 5 6 7 8 0 10 1 1 12

Total time observed (hr & min) 3:51 3 sU6 3:U6 6:38 6:35 1 0 : 5 2 6:53 2:1*5 1*:1*6 3:12 3:31 8:29 Total visits/hr. 9 h.h2 3.18 3.U5 U.37 1*.1*1 U.32 U.07 U.oo 6.92 a.0 6 a .55 7.08 Feeding only visits/hr. 9 0.26 0.27 0.27 1.06 2.13 1.38 1.71* 1.09 3.56 2.81 3.69 5.90 Brooding only visits/hr. 9 l.Oii 1.33 0.27 0.1i5 0.30 0.18 0.15 0.36 0.21 0.00 0.00 0.12 Feed and brood visits/hr■. 9 3.12 1.60 2.92 2.87 1.67 2.58 2.18 2.55 2.31 1.25 0.28 0.83 Total feeding visits/hr. 9 3.38 1.86 3.18 3.92 3.80 3.96 3.92 3.61* 5.8? a. 0 6 3.98 6.72 Total visits/hr. d 0.00 0.00 0.27 0.15 1.06 1.56 1.60 0.36 1.05 2.19 3.13 0.59 Feeding only visits/hr. d 0.00 0.00 0.27 0.15 0.91 1.1-7 1.60 0.36 1.05 2.19 3.13 0.a7 Brooding only visits/hr. d 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Feed and brood visits/hr . d 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Total feeding visits/hr. d 0.00 0.00 0.27 o.i5 0.91 1.1*7 1.60 0.36 1.05 2.19 3.13 o.a7 Total visits/hr. 9 d U.JU2 3.18 3.71 ii.52 5.1*7 5.89 5.67 1*.36 7.97 6.25 7.67 7.67 Feeding only visits/hr. 9 d 0.26 0.27 0.53 1.21 3.01* 2.85 3.31* 1.1*5 l*.6l 5 . 0 0 6.82 6.37 Brooding only visits/hr. 9 tf l.Oii 1.33 0.27 0.1*5 0.30 0.18 0.15 0.36 0.21 0.00 0.00 0.12 Feed and brood visits/hr . 90- 3.12 1.60 2.92 2.87 1.67 2.58 2.18 2.55 2.31 1.25 0.28 0.83 Total feeding visits/hr. 9 d 3.38 1.86 3.1*5 1*. 07 1*. 71 5.1*3 5.52 li.OO 6.92 6.25 7.11 7.19 602 TABLE 39

Duration of care of nestlings and nest sanitation by adult Common Grackles (Quiscalus quiscula versicolor) at West 8 - 6 1 ^ Put-in-Baya Ohio.

Days of Nest Life

1 2 3 k 5 6 7 8 9 10 11 12

Ave. duration of feed & brood visits (min) 9* 6.50 9.32 7.21 5.61 5.5k 6.00 5.33 1.00 0.00 0.00 0.00 0.00 Ave, time away from nest (min) 9. 8.1-3 8.7k 10.21 9.kl Ik.73 12. kl 13.28 10.7k 13.95 5.90 10.23 12.11 Ave, duration of feeding only visits (sec) cf. 0.00 60.38 69.09 k2.50 k0.77 55.00 1 _ Ave. duration of feed & brood visits (min) cf. 0.00 0.00 9.33 0.00 10.50 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Ave. time away from nest (min) cf. 3:3h 16.33 lk.67 13.33 15.61 12.13 lk. 56 9.56 9.93 9.00 10.86 15.56 Percent 9 cf visits with fecal sac emission. 0.6 7 0.70 0.k8 0.k9 0.60 0.65 0.67 0.67 0.85 0.51 o.56 o.ko Percent 9 cf visits with fecal sac eaten. 0.67 0.69 0.55 0.27 0.32 0.29 0.51 0.30 0.26 0.19 0.10 0.05 Percent fecal sacs eaten or carried away by 9. 1.00 0.30 0.27 0.35 0.53 0.55 0.57 0.35 0.38 0.53 o.ki 0.25 Percent fecal sacs eaten or carried away by cf. 0.00 0.70 0.73 0.65 0.57 0.55 0.53 0.65 0.62 0.57 0.59 0.75 Total time observed (hrs. & min.) 3 Ok 5:55 k:Qk 8:39 7:30 5:35 5:5 6 7:37 5:12 2 :56 5:06 2:19 210 TABLE 1*0

Duration of care of nestlings and nest sanitation by adult Common Grackles (Quiscalus quiscula versicolor) at Nest 20-61*, Put-in-Bay, Ohio.

Days of Nest Life

1 2 3 k 5 6 7 8 9 10 11 12

Ave. duration of feed & brood in I—1 c*- o visits (min) 9. 5.80 11.82 7.25 7.2k 6.77 11.87 7.80 8.25 8.83 8.66 0.00 • Ave. time away from nest (min) 9. 9.53 9.50 10.62 9.00 11.00 6.11* 11.22 9.00 7.11* 12.1*5 11.29 5.1-6 Ave. duration of feed & brood visits (min) cf. 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 Ave. time away from nest (min) cf. (or hrs. & min.) 3:51 3:1*6 3:1*5 6:37 23.20 37.07 30.25 2:1ik 38.00 23.00 16.78 31*. 00 Percent 9 cf visits with fecal sac emission. 0.1*1 0.33 0.57 0.80 0.39 0.31 0.31 0.75 0.50 1.00 0.59 o.55 Percent 9 cf visits with fecal sac eaten. o.ki 0.25 0.57 0.60 0.19 0.20 o.i5 0.1*2 0.26 0.35 0.15 0.09 Percent fecal sacs eaten or carried away by 9. 1.00 1.00 0.88 1.00 0.93 0.65 0.66 1.00 O .89 0.50 0.50 0 .86 Percent fecal sacs eaten or carried away by cf. 0.00 0.00 0.12 0.00 0.07 0.35 0.33 0.00 0.11 o.5o o.5o 0.11* Total time observed (hrs. & min.) 3:51 3 :1*6 3 :1*6 6:38 6:35 10:52 6:53 2 :1*5 1*:1*6 3:12 3:31 8:29 211 Bibliography

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