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Behavior and Taxonomic Status of Grayson's Dove 1

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Behavior and Taxonomic Status of Grayson's Dove 1 BEHAVIOR AND TAXONOMIC STATUS OF GRAYSON'S DOVE 1 LuIS 1•. BAPTISTA,2 WILLIAM I. BOARMAN, 2'3 and PARASKEVAS KANDIANIDIS 4 2Departmentof Ornithology/ Mammalogy, California Academy of Sciences, SanFrancisco, California 94118 USA; *Departmentof BiologicalSciences, San Francisco State University, SanFrancisco, California 94132 USA; and 41595San Nicolas, Ventura, California 93301 USA ABSTRACT.--Grayson'sDove (Zenaidagraysoni) is extinct in the wild but is still found in aviaries. It has been treated variously as a full speciesor as a subspeciesof the Mourning Dove (Z. macroura).The two taxa differ in shape, color, and color pattern of the rectricesand can be readily distinguishedby characteristicsof museum skins. They also differ in vocal- izationsand visualdisplays. Previous studies described differences in serology.The two taxa interbreed only rarely in captivity. It is proposedthat Z. graysonibe recognizedas a full species,closely related to Z. macroura.Received 8 November1982, accepted 8 July1983. THE Socorroor Grayson'sDove (Zenaidagray- tape recordersand analyzedon the Kay ElectricSound soni)is probably extinct in the wild (Jehl and Spectrograph($onagraph) Machine (Model 7029A) Parkes 1982). It was endemic to Socorro Island, with a 300 Hz bandwidth filter. Measurements of fre- the largest member of the Revillagigedo Ar- quency in kHz were made from wide-band sound chipelago,320 km southwestof the tip of the spectrograms.The Kay Electricamplitude display at- tachment(Model 6076C)was usedto analyzeampli- BajaCalifornia peninsula, and has been treated tude modulations within some of the vocalizations. variouslyas a full species(Friedmann et al. 1950, We measuredvarious charactersfrom study skins Blake 1953, Petersonand Chalif 1973, Ridgway after the methods outlined by Baldwin et al. (1931). 1916) or a geographic race of the Mourning The characters, followed by their abbreviations in Dove (Z. macroura)(Goodwin 1967). Short (in parentheses,are: wing length (WL), length of rectrix Mayr and Short 1970) considersit closely re- 1 (TL1), length of rectrix 2 (TL2), length of rectrix 7 lated to and possiblyconspecific with the Eared (TL7), differencebetween first- and second-longest Dove (Z. auriculata)and Mourning Dove. We rectrices(TL1-2), width of rectrixseven at widest point describeherein the vocalizationsand displays (TW), length of longest undertail covert (CL), dis- of captive Grayson'sDoves and comparethem tance from tip of longestundertail covert to tip of longestrectrix (CT-1), chordof exposedculmen (CU- with those of wild Mourning Doves. We also LEX), bill length from edge of nostril to tip of culmen discuss some external characters in the two (CULN), culmen width from baseof exposedculmen speciesand comment on the taxonomic status (CULW), and tarsuslength (TRL). of Z. graysoniand its relationship to Z. auricu- The following ratioswere calculatedfrom the above lata. measurements: WL/TL1, WL/CULN, WL/TRL, TL1/ CULN, TL1/TRL, TL1/CL, TL2/CL, TL1/TW, TL1/ METHODS AND MATERIALS TL2, CULN / TRL, CULN / CULW, TL1/TL7, TL2 / TL7, TL7 / CL. We studied the behavior of 5 Z. graysoniin indoor Analyseswere performedon the SanFrancisco State cages(3 males,2 females)and 14 in outdooraviaries. University Data Center'sCYBER 174 computer using Free-living Z. macrourawere studiedin LosAngeles, the StatisticalPackage for the SocialSciences (SPSS) Lompoc, Sonoma, and San Francisco, California. (Nie et al. 1975). Vocalizationswere taped on Nagra E and Nagra 4.2 All mensuralcharacters were log transformedand subjectedto a direct discriminant function analysis. All SPSS default criteria were utilized. Worn or dam- aged specimenswere omitted from appropriate anal- • The Grayson'sDove is also known as the Socorro ysesso that sample sizes differed depending upon Dove (see Jehl, J. R., Jr., and K. C. Parkes, 1983. "Re- variables utilized. placements" of landbird specieson Socorro Island, Materials representing seven subspeciesof Z. au- Mexico. Auk 100: 551-559). riculatawere pooled in our analyses,as we were pri- 907 The Auk 100: 907-919. October 1983 908 BAPTISTA,BOARMAN, AND KANDIANIDIS [Auk, Vol. 100 TABLE1. Descriptivestatistics on duration and other featuresof advertisingcoos and their component syllables.Syllables are labelled as in Fig. 2C, D. Z. graysoni Z. macroura Significance n œ SD Range n œ SD Range level Syllables• A 10 1.06 0.10 0.98-1.13 10 1.12 0.10 1.05-1.20 0.165 B 10 0.67 0.10 0.60-0.74 10 0.67 0.08 0.62-0.73 0.836 C 10 0.30 0.07 0.25-0.34 10 0.41 0.13 0.32-0.50 0.026 D 10 0.87 0.03 0.85-0.89 10 0.42 0.09 0.35-0.48 0.000 E 10 0.97 0.02 0.96-0.99 ..... Intersyllable intervalsa A-B 10 0.24 0.01 0.22-0.25 10 0.25 0.06 0.20-0.29 0.603 B-C 10 0.37 0.05 0.33-0.40 10 0.29 0.08 0.24-0.35 0.024 C-D 10 0.17 0.03 0.15-0.20 10 0.30 0.08 0.24-0.36 0.001 D-E 10 0.59 0.06 0.55-0.63 ..... Total duration b 10 5.24 0.25 5.06-5.42 10 3.46 0.30 3.25-3.67 0.000 Frequencyb High i0 600 32.3 577-623 9 723 31.7 699-747 0.000 Low 10 356 30.2 335-378 9 438 0 -- 0.000 Duration in seconds. Frequency in Hz. marily interested in the relationship of Z. graysonito The vocal repertoire of Zenaidadoves con- Z. macroura.The subspeciesof Z. macrourawere not sists of about six distinct utterances. These pooled, becausetwo subspecies,Z, m. macrouraand vocalizationsappear to be discreterather than Z. m. clarionensis,exhibited morphologicalcharacters graded. The first of these calls treated herein, reminiscent of Z. graysoni.We felt it necessaryto dis- the advertising coo (= perch coo of some other tinguish between charactersindicating speciesdis- investigators),is functionally equivalentto ad- tinctnessversus characterstypical of birds adapted vertising song in passerines.Jackson and Bas- for island living. Due to small sample sizesin some taxa, sexeswere kett (1964) noted that unmated males gave this pooled in all taxa. Becausemost groups show some call much more frequently than mated males. amount of sexualdimorphism, all measurementswere Thesecalls are alsogiven during aggressiveen- then subjectedto a one-way analysisof variance to counters. The second call, the nest coo, is usu- determine whether or not this would obscure the ally given by the male from the nestor the site results. There was greater variation between than where the nest is most likely to be built. The within taxa tested (P -< 0.001) for all variables. Re- third call, the greeting call, is usuallygiven by suits of discriminant analysis performed only on the male on the approach of his mate after a maleswere comparableto thoseonly for femalesand period of separation.The fourth call, the nest for both sexesgrouped. For these reasonswe feel defencecall, may be given by both sexeswhile justifiedin lumping the sexesfor all mensuralanal- ysesherein reported. Specimensexamined are listed trying to drive away an intruder from the nest. in the Appendix. The fifth call, the alarm call, is given by either sex during fearful situations.The last call, the RESULTS "growl," appearsto be given mostlyby females while calling their mates to the nest. All the VOCALIZATIONS above calls were sometimesgiven in some oth- er contexts, as discussed below. Gifford (1927) did not find any significant Advertisingcoo.--This vocalization in Zenaida differences between calls of Z. graysoniand Z. doves is homologousto the bow-coo of other macroura.Goodwin (1967) interpreted Gifford's columbiforms. Goodwin (1967) describes the statementas meaning that the calls of the two advertising coo of Z. macrouraas a disyllabic taxa do not differ. This is contrary to our ex- coo followed by two or three louder coos,or perience,as the following analyseswill show. "Coo-oo, OO, OO, OO!" We noted that some October1983] Zenaidagraysoni 909 kHz 0.5- 0 I 0 see 1 Fig. 1. Ink tracingsof somedove vocalizations.A. Advertisingcoo of Grayson'sDove. B. Advertisingcoo of Mourning Dove. C. Nest coo of Grayson'sDove. D. Nest coo of Mourning Dove. E. Nest-defencecall of Grayson'sDove. F. Greeting call of Grayson'sDove. G. Greeting call of Mourning Dove. H. Alarm call of Grayson'sDove. J. Alarm call of Mourning Dove. individuals of this specieswould add a fourth C and D is longer in Z. macroura(P -< 0.001). coo to the dissyllabicintroduction. The adver- Both highest and lowest frequenciesof the ad- tising cooof Z. graysonibegins with a disyllabic vertisingcall are higher in the Mourning Dove coo, is followed by three single coos,and ends (Table 1). with another disyllabic coo, or "Coo-oo, There are also differencesin frequencyand OO,OO,OO, Coo-oo!" This vocalization is amplitude modulation in calls of the two longer in duration than that of Z. macroura(5.24 species.Syllable A in the Mourning Dove'scall s vs. 3.46 s, Table 1). rises noticeably in frequency but does not The duration of individual syllablesand the changeappreciably in amplitude (Figs. 1, 2). intervals between them provide rhythms, Syllable A in the Grayson'scall changesonly which may function in speciesrecognition. slightly in frequency but rises noticeably in There are no differences in the durations of amplitudeabout two-thirds of the way into the syllables A and B in the two taxa. Syllable C, call. Elements B, C, and D in the Mourning however, is slightly longer in the Mourning Dove's call begin at a high amplitude and de- Dove (Table 1, two-tailed t, P -< 0.026).Syllable scendto a lower amplitude.Amplitude change D is longer in Grayson'sDove. Syllable E is in elements B and D are less noticeable in the rarely heardin Z.
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