38 WESTERN FOUNDATION OF VERTEBRATE ZOOLOGY Monograph No. 1

LITERATURE CITED (excepting items cited by title in text) DAVIS J. 1959. The Sierra Madrean element of the avifauna of the Cape District, Baja California. Condor, 61:75-84. ELBEL, R. E., and EMERSON, K. C. 1959. The taxonomic position of an Asiatic species of Otus (Ayes: Strigiformes) as indicated by the Mallophaga. Proc. Okla. Acad. Sci., 39:76-78: 1960. A new species of Kurodaia (Mallophaga: Amblycera) from the collared scops owl of Thailand. Proc. Biol. Soc. Wash., 73:119122. EMERSON, K. C. 1955. A new mallophagan from the screech-owl (Philopteridae). Proc. Ent. Soc. Wash., 57:241-242. EMERSON, K. C. and ELBEL, R. E. 1957. New species and records of Strigiphilus (Philopteridae: Mallophaga) from Thailand. Proc. Biol. Soc. Wash., 70:198-200. GRINNELL, J. 1928. A new race of screech owl from California. Auk, 45:213-214. HASBROUCK, E. M. 1893. Evolution and dichromatism in the genus Megascops. Am. Nat., 27:521-533, 638-649. JACOT, E. C. 1931. Notes on the spotted and flammulated screech owls in Arizona. Condor, 33:8-11. KNORR, 0. A. 1959. The birds of El Paso County, Colorado. Univ. Colo. Studies, Series in Biology, 5:1-48. MARSHALL, J. T., JR. 1939. Territorial behavior of the flammulated screech owl. Condor, 41:71-78. 1942. Food and habits of the spotted owl. Condor, 44:66-67. 1948. Ecologic races of song sparrows in the San Francisco Bay Region. Condor, 50:193-215, 233-256. 1957. Birds of pine-oak woodland in southern Arizona and adjacent Mexico. Pac. Coast Avifauna, 32:1-125. 1964a. (Review) The song sparrows of the Mexican plateau, by R. W. Dickerman. Auk, 81:448-451. 1964b. Voice in communication and relationships among brown towhees. Condor, 66:345-356. MILLER, A. H. 1947. The structural basis of the voice of the flammulated owl. Auk, 64:133-135. 1955. The avifauna of the Sierra del Carmen of Coahuila, Mexico. Condor, 57:154-178. MILLER, A. H. and MILLER, L. 1951. Geographic variation of the screech owls of the deserts of western North America. Condor, 53:154-178. MOORE, R. T., and MARSHALL, J. T., JR. 1959. A new race of screech owl from Oaxaca. Condor, 61:224-225. OWEN, D. F. 1963a. Variation in North American screech owls and the subspecies concept. Systematic Zool., 12:8-14. 1963b. Polymorphism in the screech owl, Otus asio, in eastern North America. Wilson Bull., 75:183-190. PHILLIPS, A. J. 1942. Notes on the migrations of the elf and flammulated screech owls. Wilson Bull., 54:132-137. SHORT, L. L. JR. 1965. Hybridization in the flickers (Colaptes) of North America. Bull. Am. Mus. Nat? Hist., 129:307-428. SLUD, P. 1964. The birds of Costa Rica. Distribution and ecology. Bull. Am. Mus. Nat. Hist., 128:1-430. STRECKER, R. L., MARSHALL, J. T., JACKSON, W. B., BARBEHENN, K. R., and JOHNSON, D. H. 1962. Pacific island rat ecology. Bull. Bishop Mus., 225:1-274. com e cbt. u-S

• • • • • • • • • •

-----;70 \ ,...... :.- *:-=':------'---

• • • • • • • • %. • • s

as;

bar bar4S

C. Qr1.Cip

Fig. 1. Author's sketches of the right foot of seven species of screech-owls, natural size, to show differences in feathering and proportions.

39 Card on evxs is

ti yra Ylt.y‘s

X0. vljf-US1 V I Yla.Va

Fig. 2. Otus asio: dorsal and ventral patterns of races within the Kennicottii and Seductus groups, dorsal ground color indicated.

40 ocdli

lot of C1r)riq.1"NOK

yvv.xx We.

0

Se»310)e; QS I0

4:ficereAni- cc, r fro K., bc.ck cleo.rer 2 less row.% or IOv ft-) darker htbrt. tse Iy pn.o.rkedi 6t4,k.

Fig. 3. Otus asio: color patterns of the Cooperi and Asio groups.

41 n1PSameri LQYtU5

n tom h'rti C0.SStni s httus Ua ha oia e, ciorsaii fOni -Hnovn Sons II cictcr isicfc..,tus are. fhe Sawte.

LA a•t-LLS 0.1m len ta. 01 US (venter dacr SISI-Cktfu

Fig. 4. Otus triehopsis, 0. guatemalue, and 0. flanneolus: dorsal and ventral black patterns with dorsal ground color indicated.

42 NV,4

0 b./5 C to rk tarsus fec....{-4e (el ower tarsus n.ke4 extr toffs lo.r e, ear 4-4-1.5 rtserti- buck not aisfirtc.f;v y 01co rej

LAJ 0

615er

4F--- bcxY t9car u ( ScAle t 45, -h-; lavAyueoios f svi.o.iter foot-

4,4Lete ft.0.1-kken ev.ei

ot vs c 1, I., rk) 0- +arts rtake.4 ecortv f ts re.Sert 3

Fig. 5. Otus barbarus, 0. clarkii, and O. choliba: dorsal and ventral coloration.

43 44 WESTERN FOUNDATION OF VERTEBRATE ZOOLOGY Monograph No. 1

EXPLANATION OF FIGURE 6 (opposite page):

Subspecies names are placed in the approximate region where each race attains its greatest distinctiveness. Because of a higher stand- ard than the conventional one for subspecies, my 100% distinguishable races involve two awkward shifts. I synonymize macfarlanei into bendirei by the rules of priority but show the best development of bendirei in the geographic range of macfarlanei. Similarly, I show the best development of asio in the territory of its synonym, naevius. This is because the older name's type specimen comes from an area of interme- diacy. On the other hand there are intermediate races whose type locality is exactly in the proper intermediate area. Some are well-known and venerable such as cineraceus, inyoensis, and swenki.

For those museum curators who prefer to retain all these valid, but decidedly intermediate subspecies, the table of synonyms in the appendix shows the correct names, and their distributions appear as "intermediate races" in the sections on intergrading in the appropriate subspecies accounts.

In summary, my disposition of well-known names from the AOU Check-list is as follows: The intermediate race asio of the Check-list has its name replacing naerius to become the name of a distinctive subspecies. The intermediate race benclirei of the Check-list has its name replacing macfarlanei to become the name of a distinctive subspecies. The intermediate race cineraceus of the Check-list is submerged into aikeni. The intermediate race swenki of the Check-list is submerged into maxwelliae.

70 80 84 84 80 70

: ":', , P 4V4740111r, 4,,,. 1 50 W, 180 * 170 0 '0 Villikti ,.ffr"

1111144111* ° 60

k`R it 'AA.

gi 111111°'

.8 ...,

L., : If, .. ''.,.N . \ Piie) -s1-, 50 --. .___. .... „oU4

1

e=> I " i kennicotti i Iii i •r. 4g 't v APP L. A \ 1 4

40 ,_.... .,) ' 40 • 1I \ maxwell ice ._._) bendirei asio . 7. — ---/-1 , .- .1--• .1. _I _._. 1. is, KENNICOTTII ASIO (___.-- - V' \ r----- _,..-. DO aikeni i r 111111F yuman- • I so •-ensis i -•,... .."-• •••____ i --- - i .t..---- hasbroucki I cardonensis 7: \ it • - floridanus vin- 0 .0.---4. aceui • mccallil '`i

Silt— i..... 20 xantusi toni d 20 r

Mitik \ I eZ=L h .- L. „_, .7T-- ..-1( , SEDUCTUS . , Sr" ) I / '-- Ofus osia Iambi /

Museum of Vertebrate Z.. ology COOPERI % 0 1„q3 3 cooper -,- ale a 0 100 208 300 900 500 8caleofMiles !bli 1111

130 120 110 IOD 90 80 70

Fig. 6. Otus asio: distribution of the four race groups.

45 46 WESTERN FOUNDATION OF VERTEBRATE ZOOLOGY Monograph No. 1

EXPLANATION OF FIGURE 7 (opposite page):

Locality Museum Date Phenotype (if not worn) KENNICOTTII GROUP, SOLID CIRCLES Pine Springs, Culberson County, Denver Sept 1945 8 9 aikeni and 8 mi. N Guadalupe Peak MVZ March 1950 aikeni Bell Canyon (cannot find) "70 mi. Peabody June 1947 8 suttoni NE Van Horn, 5000 ft." Lajitas Arizona, Oklahoma Oct 1961 3 a 19 suttoni Terlingua USNM June 1901 suttoni and 2 juveniles Chisos Mts Univ. Texas Sept 1937 Big Bend Park July 1962 8 .suttoni Boquillas Arizona July 1962 8 suttoni of mixed pair, another lone 8 heard 9 mi. S Boquillas Chicago April 1945 8 of aikeni phenotype Sierra del Carmen USNM, MVZ Aug 1936, 5 a 4 9 suttoni April 1953 Langtry Arizona Oct 1961 18 19 suttoni 1 8 suttoni 2 mi. west seen Juno USNM July 1902 9 suttoni and juvenile

ASIO GROUP, TRIANGLES Sheffield July 1962 8 heard and seen Independence Creek U. Texas, Arizona July 1949, 2 8 2 9 , others heard July 1962 Boquillas Arizona July 1962 1 ,8 juvenile mccallii and 1 9 mccallii of mixed pair 10 mi. W Langtry Arizona July 1962 1$ 19 Langtry Arizona Oct 1961 3 S : mccallii, cineraceus and hasbrottcki Comstock USNM June 1902 y mccallii mouth of Devil's River, and Del Kansas, MVZ Feb 1953 2 9 mccallii Rio, 4 mi. N. Dec 1954 Fort Clark USNM March 1893, 19 of hasbroucki phenotype April and May 1898 1 8 and 1 9 mccallii Jimenez Kansas June 1952 18 mccallii Nave July 1962 18 heard Allende, 8 mil. S. Arizona July 1962 19 mccallii La Gacha Kansas Dec 1953 1 mccallii La Mariposa U. Texas March 1938 8 mccallii

ASIO GROUP I

• Pine Springs and I near Guadalupe Pk.

KENNICOTTI I GROUP

O ? Bell Canyon near Sheffield "P • • Independence Creek

O Juno w of Langtry

Langtry Comstock Terlingua mouth Devil's R. k•ABoquillas • Ft. Clark Lajitas • • 41rear. " • Jime nez Chisos Mts. • Sa del Carmen • Nava • La Mariposa • near Allende La Gacha

0 50 100 miles

Otus asio

Fig. 7. Otus asio: overlap of Kennicottii and Asio groups at the Big Bend of the Rio Grande. 47 OTUS TRICHOPSIS

...: PHENOTYPE o o o DORSAL / VENTRAL COLORATION ..o / 6, c" i • :, +. .' '. so ,.. co -4* ,ri' 0Q c0....\ 0- 4..-- cf ,i.i .6 ° Au .o q o o ..9 P° o

Arizona, Sonora and Chihuahua

Nuevo Lean (tentative), Zacatecas, San Luis Potosi

ri—Lsi— solid = red phase

Sinaloa, Jalisco, Nayarit and Colima type of ridgwayi

Michoacan, D. F. and Veracruz

H Guerrero, Oaxaca type of enano cotype of enano

Chiapas and Guatemala

1T-1 NW El Salvador type of mesamericanus type of pumilus

NE El Salvador, Honduras and Nicaragua

Fig. 8. Otus trichopsis: coloration of 129 fall and winter specimens. (Each small square represents one museum specimen. All specimens in a vertical column are of the same dorsal/ventral coloration, named at the top of that column; all specimens in a horizontal array are from the same population. Red phase is blacked in. For brevity, the name "enano" is used for the phenotype predominant in the Chiapas-Guatemala population.)

48

OT US FL AMMEOLUS

co

PHENOTYPES .c co oco 0

€ POPULATIONS

Pacific sta tes and

Idaho, idahoensis 11=1•1•111101111•1•101111•• •=1

Great Basin and southern Rockies AL.

Southern border of USA, northern Mexico, flammeolus ?

Mexico south of. Tropic of Cancer, rarus

Fig. 9. Otus flammeolus: coloration of 32 fall-plumaged birds thought to be on their breeding grounds. 49 50

Fig. 10. Diagrammatic representations of sonograms of Screech-Owl songs .Otus— asio and Otus trichopsis. (Otus cooperi was recorded by Irby Davis. The opening trill of its primary song is not frequently used; it and the first trill of the double trill are confused on the sonograph, and do not agree with what the ear discerns, especially when the recording is played slowly. The sonogram is reproduced here mainly because it is the only known recording of this owl; it is atypical and unfortunately does not support the author’s claim that cooperi is conspecific with asio, based on his field notes and diagrams of normal songs of cooperi males. The rest of the owls were all recorded by the author.) 51

Fig. 11. Diagrammatic representations of sonograms of Screech-Owl songs. (Otus guatemalae was recorded by Irby Davis, O. choliba by Paul Schwartz, and O. scops by Tsuruhiko Kabaya. The rest were recorded by the author.) Bob Broder Fig. 12. Otus asio, Kennicottii Group: lower, kennicottii, coastal British Columbia; upper, bendirei, interior Oregon. 52 Bob Broder

Fig. 13. Otus asio, Kennicottii Group: ventral view of same specimens shown in Fig. 12.

53 Bob Broder Fig. 14. Olus usio, Kennicottii Group: lower, tending toward cardonensis phenotype but ventral pattern insufficiently dense (see Fig. 15), San Diego County, California; center, suttoni, Mexico, D. F.; upper, xantusi, Baja California Sur (with ventral cross-bars atypically vague—see Fig. 15).

54 Bob Broder

Fig. 15. Otus asio, Kennicottii Group: ventral view of same specimens shown in Fig. 14. 55 Bob Broder

Fig. 16. Otus asio, Kennicottii Group: lower, aikeni, Mogollon Plateau, Arizona; center, cineraceus, Fort iluachuca, Arizona; upper, yurnanensis, Yuma Arizona. 56 Bob Broder

Fig. 17. Otus asio, Kennicottii Group: ventral view of same specimens shown in Fig. 16. Observe that cineraceus is intermediate in all respects between yumanensis and aikeni. 57 Bob Broder

Fig. 18. The vinaceous screech-owls of the Pacific coast of Mexico. Otus asio, Kennicottii, Seductus, and Cooperi groups: bottom to top—vinaceus, coastal Sinaloa; seductus, Colima; Iambi, Oaxaca.

58 Bob Broder

Fig. 19. The vinaceous screech-owls of the Pacific coast of Mexico—ventral view of the same specimens shown in Fig. 18.

59 Bob Broder

Fig. 20. Otus asio, Cooperi and Asio groups: bottom to top—cooperi, Costa Rica; maxwelliae, Colorado; asio, Iowa. 60 Bob Broder

Fig. 21. Ventral view of same specimens shown in Fig. 20. (Note: All figures of plumage are of the normal phase; none is of the red phase. either in the drawings or in the photographs.) 61 Bob Broder

Fig. 22. &us asio, Asio Group: top to bottom—mccallii, Langtry, Texas; hasbroucki, Kerrville, Texas; floriclanus, Florida. 62 Bob Broder Fig. 23. Otus asio, Asio Group: ventral view of same specimens shown in Fig. 22. The upper specimen, mccallii, has atypical coarse ventral markings see Fig. 31).

63 Bob Broder

24. Olus asio and Otus trichopsis:. bottom to top—Otus asio semplei, San Luis Potosi; Olus trichopsis aspersus, Arizona; Outs trichopsis ssp.?, San Luis Potosi; Olus 1. trichopsis, Mexico, D. F.

64 Bob Broder Fig. 25. Otus asio and 0. trichopsis: ventral view of same specimens shown in Fig. 24. 65 Bob Broder Fig. 26. Otus trichopsis and Otus guatemalue: bottom to top—Otus trichopsis ssp.?. Chiapas; Ones trichopsis mesamericanus, Mt. Cacaguatique, El Salvador; Otus guatemalae liastatus, Sonora; Otus guutemalae cassini, Tamaulipas.

66 Bob Broder

Fig. 27. Otus trichopsis and 0. guatentulue: ventral view of same specimens shown in Fig. 26. 67 Bob Broder

Fig. 28. Otus guatemalae: bottom to top—thompsotii, Yucatan; guatemalae, Honduras; black guatemalae, Guatemala; dacrysistactus, Nicaragua. (Note: Even though I do not recognize all races, I use their names for discernible phenotypes.)

68 Bob 13roder

Fig. 29. Otus guatemalae: ventral view of same specimens shown in Fig. 28. 69 ,,AC 7 re., ft•k• • (.7.,VL

Bob Broder

Fig. 30. Otus guatemalae and Otus flammeolus: bottom to top—Otus guatemalae vermiculatus, Panama; Otus flammeohts, black Great Basin-southern Rocky Mts. phenotype, Hualpai Mts., Arizona; 0. flammeolus, typical phenotype, Catalina Mts., Arizona; 0. flammeolus, red rants phenotype, Davis, California. 70 Bob Bioder

Fig. 31. Ottis guatemalne and 0. flammeolus: ventral view of same specimens shown in Fig. 30. 71

Monograph No. 1 72 WESTERN FOUNDATION OF VERTEBRATE ZOOLOGY ABBREVIATED TOPICAL INDEX

Page: Topic: Page: Topic:

26 Acknowledgements 31 cassinii, Otus guatemalae

17 Appendix 32 cooperi, Otus asio

21 Comments on type specimens 34 floridanus, Otus asio

26 Concealing coloration and predation 1 guatemalae, Otus guatemalae

19 Conclusions 27 hasbroucki, Otus asio

hastatus, Otus guatemalae 26

Geographic variation: kennic•ottii, Otus asio 12

Otus asio 5 Iambi, Otus asio 17

Otus guatemalae 25 maxwelliae, Otus asio 19

Otus trichopsis 22 mccallii, Otus asio 20

The Asio Group 18 mesamericanus, Otus guatemalae 23

The Cooperi Group 17 seduc•tus, Otus asio 16

The Kennicottii Group 11 suttoni, Otus asio 14

(Introduction) 1 trichopsis, Otus trichopsis 23

vermiculatus, Otus guatemalae 26

Incipient species: vinaceus, Otus asio 15

Asio Group, Eastern screech-owl 18 xantusi, Otus asio 15

Cooperi Group, Pacific screech-owl 16 yurnanensis, Otus asio 14 Kennicottii Group, Western screech-owl 8 Red Phase specimens (Kennicottii Group 35 Seductus Group, Balsas screech-owl 16 Key to species and incipient species 4 Species accounts: Literature cited 38 Otus asio, common screech-owl 6 Museums 36 Otus barbarus, bearded screech-owl 27 Parallelism and natural selection 2 Otus c•holiba, tropical screech-owl 24 Otus clarkii, bare-shanked screech-owl 27 Recognized subspecies: Otus flammeolus, flammulated owl 23 aikeni, Otus asio 13 Otus guatemalae, vermiculated screech-owl 25 asio, Otus• asio 20 Otus trichopsis, whiskered screech-owl 21 aspersus, Otus trichopsis 23 Synonyms 33 betzdirei, Otus asio 12 Synopsis 32 cardonensis, Otus asio 13 The species question ... 3