418 SHORT COMMUNICATIONS

The Condor98:418-422 0 The CooperOrnithological Society 1996

NUPTIAL VOCALIZATIONS OF MALE LEAST SEEDSNIPE: STRUCTURE AND EVOLUTIONARY SIGNIFICANCE’

EDWARDH. MILLER Biology Department, Memorial Universityof Newfoundland,St. Johns,’ NF AlB 3X9, Canada, e-mail: [email protected]

Key words: Thinocorus rumicivorus, Least Seed- METHODS snipe;vocalization; systematics. Least Seedsnipewere recordedopportunistically about 85 km NW of Rio Gallegos,Argentina, in November Acoustic displays of non-passerineshold promise for 1977. Recordings of Common Snipe (G. g. delicata) understandinghow communication is adapted to the from severalNorth American locationsincluding Alas- physical environment and for resolving systematicre- ka, Yukon, British Columbia, and Manitoba were an- lationships.The potential ofacoustic characteristicsfor alyzed for comparative purposes.Recordings of Com- systematicscan be illustrated with shorebirds. Some mon Snipe (G. g. gallinago) from Iceland and Russia shorebirddisplays (e.g., snipe drumming) differ greatly were also analyzed(Russian recordings were taken from between closely related species, so should be infor- Veprintsev [ 19821).All recordingsexcept Veprintsev’s mative about low-level relationships. Other displays were made at 19 cm/set with a Nagra IS tape recorder (e.g., duetting in stone-curlews, avocets, and oyster- and Scotch 208 tape, with a Sennheiser MKH 816 catchers;Stepanjan 1970, 1979)are evolutionarily con- “shotgun” microphone. Analyses were carried out on servative and extremely similar even between shore- a microcomputer with the signal-analysispackage CSL bird speciesthat diverged from one another millions 4300 (Ray Elemetrics Co., Pine Brook, New Jersey, of years ago; such conservative displays may help to 07058). Sound samples were digitized at 20,000 Hz. resolve high-order relationships. For the sonagramsshown below, analyseswere done A shorebird group whose systematic position may with a Blackmanwindow, no pre-emphasis,and frame be clarified by acoustic information is the seedsnipe sizescorresponding to an analoganalyzing filter band- (Thinocoridae). The relationships of the four species width of 50-60 Hz. Descriptions and measurements of seedsnipeto other shorebirdsremain enigmatic de- given below are based on six to ten examples of each spite a wealth of systematic and phylogenetic studies kind of vocalization, from an estimated 1O-l 5 indi- over many years (Strauch 1978, Sibley and Ahlquist vidual birds recorded over four days. 1990, Bjiirklund 1994, Chu 1995). The purposeof this paper is to describesome nuptial vocalizationsof seed- RESULTS snipe and make adaptive and phylogeneticinferences Male seedsnipecommonly perch on prominencessuch based on acousticstructure. as a knoll, bush, or fencepost and utter two kinds of Vocalizations of seedsnipehave not been analyzed loud calls in long series. Spectrographically,the two spectrographically(Miller 1984, 1992, 1995), although some behavioral descriptions exist (Maclean 1969, call types are distinguishablefrom song elements. To Fjeldsa and Krabbe 1990). In this note, I presentspec- the ear, however, there is some similarity, which ex- trographicanalyses of long-rangevocalizations by male plains why Maclean (1969) applied the same terms to both. For example, he referred to “a rapid pu-pu-pu- Least Seedsnipe (Thinocorus rumicivorus),based on observationsduring the breeding seasonin Patagonia. pu-pu, derived from the first syllable of. . . puku” and Three kinds of vocalization are described:a complex (for the Gray-breasted Seedsnipe, T. orbignyianus) aerial vocalization and two simpler vocalizations that lengthy “songs” of “as many as 850 puku notes” while are uttered when on the ground or a prominence. Vari- perched.These descriptionslikely refer to the two types ations and other forms of ground and flight display of simple calls recognizedherein. One of these call types is a series of rhythmically occur (Maclean 1969) but my tape recordingsare not adequate for their spectrographicanalysis. These vo- repeatedsimple notes(Fig. 1A). The noteshave a mod- calizationsresemble those of many scolopacidsin their erately broad bandwidth (-0.6-1.2 kHz), though no rhythmicity, complexity,and syntacticalproperties, and harmonicsare present:most energyis at -0.9-l. 1 kHz. are notably similar to some vocalizations of Common The notes are brief (- 15-25 mseclong), with intemote Snipe (Gallinago gallinago). Despite the strong simi- intervals (INIs) of -200-220 msec. for a call rate of larities, phylogeneticconclusions cannot be drawn be- -4/set and -9% coverageby vocalizationsduring call causeof the paucity of acousticdata for most species. sequences(i.e., 9 1% silence). The secondkind of simple call is also uttered rhyth- mically, but is organizedas two alternating note types (Fig. 1B). One type spans -0.8-1.3 kHz, with most energy over - 1.0-1.3 kHz. These notes are -50-55 I Received 11 July 1995. Accepted 9 February 1996. msec long. Briefer notes (- 20-25 msec long) alternate SHORT COMMUNICATIONS 419

-80% silence). The main differences from seedsnipe are frequencymodulation and harmonic structure,the latter being responsiblefor the broad bandwidth and high frequenciesreached. The most complex vocalization of male Least Seed- snipe is aerial “song.” In the early morning, when it is still very dark, male seedsniperise silently in aerial displaysto a height of - 10 m then, just after maximal height is reached, start to glide gradually downward while uttering a long complex vocalization; toward the end of the glide they may dive before landing (Maclean 19691observed heights up to 15-20 m). Males seemed LL [ - 500 msec to give these aerial displays only a few times each morning, and sporadicallyduring the daylight hours. 3.0 c Song is a vocalization up to 15 set long consisting 2.0 offour to five note types(Fig. 3A). It beginswith several 1.0 1 ’ /r m 0 1 7 soft notesthat increasesuccessively in frequency.They are followed by a long sequenceof alternating loud note types in short series (wikiti of Maclean [ 19691). One note type spans -0.6-1.2 kHz and is very brief Harmonic structureis absent.These briefnotes (pulses) occur in triplets in Figures 3A-B, with a decline in frequency and increasein duration occurringprogres- FIGURE 1. Sonagramsof ground/perch advertise- sively over each triplet (note durations increase from ment calls by male Least Seedsnipe.A-Part of long -20 to -40 msec within triplets). The pulse triplets calling sequenceof singlenotes. B-Part of long calling alternate with one or two longer (-55-95 msec) notes sequenceof double notes. C-Part of(B), on logarith- at lower frequency (-0.6-0.9 kHz). The longer notes mic frequency scaleand different time scale.Analyses characteristicallyrise, sometimes in step-fashion,to a were done over the frequencyranges 100-2000 Hz (A- constant-frequencyportion, then drop off quickly at B) and 100-5000 Hz (C).

with them, and are -0.8-l. 1 kHz in frequency. Inter- vals are - 140-l 45 msec long following long notes and - 115-l 30 msec long following brief notes. No har- monics are presentin either note type. Call rate is - 5- 6/set, with vocalizationsaccounting for 22% of calling time (=78% silence within calls). II Common Snipe have two well known vocal forms (, 625 msec similar to thosejust describedfor the Least Seedsnipe. As in the latter species,Common Snipe also give them from the ground or prominences, including mounds, fenceposts,houses, power lines, and trees. Both call types are uttered during parts of aerial displaysas well (Miller, unpubl.). The first of the two loud call types is the Chip (often termed yak). The secondis the Chip- per (often termedjick-jack) The names Chip and Chip- per follow Cramp (1983). Sonagramsof thesecall types are in Grudzien (1976) for delicuta,and in Glutz et al. (1977), Reddig (1978, 1981), Bergmann and Helb (1982), and Cramp ( 1983) for gdinugo. The Chip is uttered rhythmically in long monoto- nous series at a rate of -2-4/set. It has two variants that may represent distinct call types, one shaped as IE - 250 msec an inverted chevron on sonagrams(Fig. 2A), the other as a chevron (Fig. 2B). They are -50-80 msec long, FIGURE 2. Sonagramsof ground/perch advertise- separated by intervals of -190-330 msec (yielding ment calls by male Common Snipe, illustrating single- -21% vocal coverage = 79% silence). The Chipper and double-call forms resemblingthose of Least Seed- likewise is uttered in long monotonous series,but at a snipe. A-Part of long calling sequenceof singlenotes higher rate (-5-6/set; Fig. 2C-E). It consists of two (G. g. delicutu).B-Ditto, for G. g. gullinugo.C-Part alternatingnote typesthat differ in frequencyattributes of long calling sequenceof double notes(G. g. delicutu). (see Fig. 1E) but are similar in temporal features to D-Ditto, for G. g. gullinugo. E-Part of C, on loga- those of the Least Seedsnipe:brief notes are -20-40 rithmic frequencyscale and different time scale.Anal- msec,long notesare -45-50 msecand INIs are - 140- yses were done over the frequency ranges 100-2000 180 msec long (vocal coverage - 17-21% of time = Hz (A-D) and 100-5000 Hz (E). 420 SHORT COMMUNICATIONS

B -200msec c -2OOmseC

FIGURE 3. Sonagramsof song in display flight by male Least Seedsnipe.A-Complete song (marked parts are shown in B and C, on logarithmic frequency scale and different time scale). Analyses were done over the frequency range 100-2000 Hz. the end. During this portion of song,notes are uttered this species are low in frequency (e.g., song is only at a rate of -9-lO/sec, and occupy -40% of a song’s - 500Hz-1.3 kHz) consideringits small size: 14 males duration. in the Royal Ontario Museum averaged 68 g in fresh A transition occursabout halfway througheach song, weight (Dunning [ 19931gives estimatesof 40-45 g). In with a switch to lower-frequency and longer notes re- this respect,the Least Seedsnipeis similar to another semblingthe long notesjust described(Fig. 3C). These small (- 60 g; Dunning 1993)open-country species that notes decline in frequency overall (to ~600 Hz at the advertises on the ground, the Plains-wanderer (Pe- end), are longer (-90-130 msec), narrower in band- dionomus torquatus),which has long-distanceadver- width, and have simpler frequency modulation than tising calls at - 15-570 Hz (mainly 300-350 Hz) (Pet- earlier notes in the song. The later notes are the “far- tigrew and Larsen 1990). carrying puku puku puku” sounds described by Ma- The simple structure and rhythmic repetition of clean (1969). The brief pulsed notes drop out, so the ground/perch vocalizations enhance the ability of lis- rate of calling declines (5-6/set), and vocalization oc- teners to accuratelyperceive the soundsover long dis- cupies more of the time (- 56% = 44% silence). tances (Schleidt 1973). More subtle information can Songof Common Snipe is a long complex utterance be conveyedover long distancesby gradualchanges in that is given at various times during Drumming display frequency, loudness, and duration that take place flights, particularly during the terminal dive. It com- throughout calling sequences,as listeners can detect prises alternating sequencesof note types similar to the trends even if some notes are degradedor parts of those illustrated in Figure 2 (Miller, unpubl.). a call sequencecannot be heard. Such sequentialgrad- ing is simplestin one-note calls;in two-note calls,grad- ADAPTATIONS IN VOCAL BEHAVIOR AND ing occursat several levels-as individual notes, note ACOUSTIC STRUCTURE pairs, and associatedINIs are affected. Effective long- distance communication through multi-level sequen- Vocal behavior ofLeast Seedsnipeshows general agree- tial gradingis also likely for song,despite its high struc- ment with predictions about adaptations for signal tural complexity and hierarchicalorganization. For ex- transmissionin open environments (Wiley and Rich- ample, at a coarselevel, the song in Figure 3 exhibits ards 1982). The use of bushesand other prominences a general decline in frequency after the first - 5 notes; as calling postsreduces acoustic degradation caused by at finer levels, the brief notes organized as triplets in interference with the ground surface and vegetation. Figure 3B show sequentialgrading, and the note group- Vocalizing in aerial display reducesdegradation sim- ings which include the triplets themselves commonly ilarly, and alsoincreases broadcast range. As well, Least exhibit sequentialgrading, although this is not appar- Seedsnipe song is most common in the very early ent in Figure 3. morning, when the high atmospheric turbulence that characterizesopen environments is minimal. Some frequency characteristicsof Least Sandpiper PHYLOGENY vocalizations likewise seem interpretable as adapta- Least Seedsnipevocalizations show strong similarities tions for long-distancetransmission in open environ- to vocalizations of Common Snipe and some calidri- ments. Unlike many open-countryspecies, Least Seed- dine sandpipers. Some similarities are in high-order snipe vocalizations are not especiallynarrow in band- attributes of vocalizations such as repertoire charac- width, but the absence of harmonic structure never- teristics,note-group structure,and sequencing.As not- theless results in the integrity of spectral information ed above, high-order attributesare predicted to evolve over relatively long distances.Low-frequency sounds most slowly, hence to be most useful for investigating attenuate least over distance, and all vocalizations of affinities between distantly related groups. SHORT COMMUNICATIONS 421

A repertoirecharacteristic shared by Least Seedsnipe amona Charadriiformes: reanalysis of previous and Common Snipe is the presenceof similar one- and data. Auk 111:825-832. _ two-part ground/perchcalls. The calls are structurally CHU. P. 1995. Phvloeenetic reanalvsis of Strauch’s similar in the brevity and simplicity oftheir constituent bsteologicaldata se; for the Charadriiformes. Con- elementsand in other features:percent of time covered dor 97:174-196. by calls; rhythmic repetition in long seriesresulting in CRAMP,S. [ed.]. 1983. Handbook of the birds of Eu- “tonic” displaying;and, for the two-note calls, a syn- rope, the Middle East and North Africa. The birds tactical arrangementof alternating note types and as- of the Western Palearctic, Vol. 3: waders to gulls. sociatedIN1 durations. The calls are also functionally Oxford Univ. Press, Oxford. similar, in being long-distancedisplays used by breed- DUNNING,J. B., JR. 1993. CRC handbook of avian ing males from the ground or perches.Considering the body masses.CRC Press, Boca Raton, FL. great phylogeneticdistance between the Thinocoridae FJELDS&J., ANDN. K~ABBE. 1990. Birds of the High and other shorebird taxa (Sibley and Ahlquist 1991, Andes. A manual to the birds of the temperate Bjiirklund 1994, Chu 1995), I hesitate to suggestthat zone of the Andes and Patagonia,South America. the two vocal types are homologous between Least Zoological Museum, Apollo Books, Svendborg, Seedsnipe and Common Snipe; however, it is note- Denmark. worthy that very similar organization to the two-part GLUTZ, U. N., K. M. BAUER,AND E. BEZZEL[eds.]. call occursin other scolopacidssuch as the Chatter call 1977. Handbuch der Vdgel Mitteleuropas, Band of Least and Baird’s Sandpipers(C. bairdii) and Mo- 7: Charadriiformes. Akademische Verlagsgesell- torboat Sound of Semipalmated Sandpiper(C. pusilla) schaft, Wiesbaden, Germany. (Miller 1983a, 1983b, Miller et al. 1988). The rapid, GRUDZIEN,T. A. 1976. The breeding behavior of the lengthy, and rhythmic repetition of a set of brief notes Common Snipe (Gallinagogallinago delicata Ord) that differ from one another and sometimes differ also in central Michigan. M.S.thesis, Central Michigan in associatedINIs (e.g., Figs IB-C, 2) may therefore Univ., Mount Pleasant, MI. be a very old feature of vocal organization in the Scol- LEMON, R. E. 1977. Bird song: an acoustic flag. opacida (= Jacanidae, Rostratulidae, Pedionomidae, Bioscience 27:402408. Scolopacidae,and Thinocoridae; Sibley and Ahlquist MACLEAN, G. L. 1969. A study of seedsnipein south- 1990). em South America. Living Bird 8:33-80. Some simple syntactical properties are progressive MILLER,E. H. 1983a. Structure of display flights in changesin loudness,frequency, and call rate over the the Least Sandpiper. Condor 85:220-242. courseof a song(Lemon 1977). More complex syntax MILLER,E. H. 1983b. The structureof aerial displays is the organization of notes into groups, nonrandom in three speciesof Calidridinae(Scolopacidae). Auk sequencingof both attributes and note types within 100:440-451. and acrossnote groups, and characteristicsof multi- MILLER, E. H. 1984. Communication in breeding level sequential grading. At this level of description, shorebirds,p. 169-24 1. In J. Burgerand B. L. Olla thesesyntactical properties all occurin Least Seedsnipe [eds.], Shorebirds: breeding behavior and popu- song,and hold promise for use in phylogeneticstudies. lations. Plenum Press, New York. At present, however, neither thesenor any other vocal MILLER, E. H. 1992. Acoustic signalsof shorebirds. charactersof Least Seedsnipeshed light on the family’s Royal B. C. Museum Tech. Rep., Victoria, British systematic position. To make progress in this area, Columbia. more detailed behavioral and spectrographicinfor- MILLER, E. H. 1995. Sounds of shorebirds: oppor- mation about nuptial vocalizationsis neededfrom oth- tunities for amateurs and an update of published er shorebird species,beginning with T. orbignyianus information. Wader Study Group Bull. 78: 18-22. and Attagis. At present, even available analysesof G. MILLER,E. H. 1996. Acousticdifferentiation and spe- gallinagovocalizations are inadequatefor phylogenetic ciation in shorebirds,p. 24 l-257. In D. E. Kroods- studies(Miller 1996). ma and E. H. Miller [eds.], Ecologyand evolution of acousticcommunication in birds. Cornell Univ. I am indebted to Allan Baker of the Royal Ontario Press, Ithaca, NY. Museum for inviting me to participatein his expedition MILLER.E. H.. W.W.H. GUNN. ANDB. N. VEPRINTSEV. to Patagoniaand for encouragingme publish theseob- 1988. Breedingvocalizations of Baird’s Sandpip- servations.That expedition was supportedby the Roy- er Calidris bairdii and related species, with re- al Ontario Museum and the Natural Sciencesand En- marks on phylogenyand adaptation. Omis Stand. gineeringResearch Council of Canada (operatinggrant 19~257-267. to AJB). Analysis and writing were supportedby Me- PETTIGREW, J. D., AND 0. N. LARSEN. 1990. Direc- morial University of Newfoundland. Brad Millen of tional hearingin the Plains-wanderer,Pedionomus the Royal Ontario Museum kindly provided infor- torquatus,p. 179-190. In M. Rowe and L. Aitkin mation on body weights. Allan Baker reviewed a [eds.], Information processingin mammalian au- manuscript draft. ditorv and tactile systems.Alan R. Liss Inc., New LITERATURE CITED York: REDDIG,E. 1978. Der Ausdrucksflugder Bekassine BERGMANN,H.-H., ANDH.-W. HELB. 1982. Stimmen (Capellagallinagogallinago). J. Omithol. 119:357- der Vogel Europas. Gesslngeund Rufe von tlber 387. 400 Vogelarten in mehr als 2000 Sonagrammen. REDDIG,E. 1981. Die Bekassine Capella gallinago. BLV Verlagsgesellschaft,Miinchen, Germany. Die Neue Brehm-Buecherei, A. Ziemsen Verlag, BJ~~RKLUND,M. 1994. Phylogenetic relationships Wittenberg Lutherstadt, Germany. 422 SHORT COMMUNICATIONS

SCHLEIDT,W. M. 1973. Tonic communication: con- STRAUCH,J. G. 1978. The phylogeny of the Char- tinual effects of discrete signsin animal commu- adriiformes (Aves): a new estimateusing the meth- nication systems.J. theor. Biol. 42:359-386. od of charactercompatibility analysis.Trans. Zool. SIBLEY, C. G., ANDJ. E. AHLQUIST. 1990. Phylogeny Sot. Lond. 34:263-345. and classificationof birds. A study in molecular VEP~UNTSEV,B. N. 1982. Birds of the Soviet Union: evolution. Yale Univ. Press. New Haven. CT. a soundguide. Three long-playingdiscs. Melodiya, STEPANYAN, L. S. 1970. (Non-morphological criteria All-Union Studio for Recorded Sound. Moscow. and their utilization for classification[Burhinidae (Narration in Russian) (Aves) taken as an example]). Z. obshch. biol. 3 1: WILEY, R. H., AND D. G. RICHARDS. 1982. Adapta- 29 l-301. (In Russian) tions for acousticcommunication in birds: sound STEPANYAN,L. S. 1979. (Possible relations of Zbi- transmissionand signaldetection, p. 13l-l 8 1. In dorhynchastruthersii and notes on the history of D. E. Kroodsma and E. H. Miller [eds.], Acoustic the family Haematopodidae). Zool. Zh. 58:167 l- communication in birds, Vol. 1. Academic Press. 1679. (In Russian) NY.

The Condor98:422-423 0 The CooperOrnithological Society 1996

FLORIDA SCRUB-JAY FORAGES ON BACK OF WHITE-TAILED DEER’

JOHNW. F~~~PA~~IcK~AND GLENE. WOOLFENDEN ArchboldBiological Station, P.O. Box 2057, Lake Placid, FL 33862-2057

Key words: Foraging; mammals; cleaningstation; is based on careful study of his video, a copy of which symbioses;tameness; Florida Scrub-Jay;Aphelocoma is depositedin the archivesat Archbold Biological Sta- coerulescens. tion. On 25 September 1994, Mr. Andrew Reiner was The propensity of Florida Scrub-Jays (Aphelocoma video-recording scenery and deer movements from a coerulescens)to become extremely tame around hu- deer-standoutside of, but immediately adjacentto. the mans, willingly to perch and even to rest on the hand, westernboundary of Archbold Biolog&alStation, near shoulder, or head, has been commented upon by nat- Lake Placid, Highlands Countv. Florida. At annroxi- uralists for over a century. We hypothesized (Wool- mately 07:Ob EDT a large buck deer with full-antlers fenden and Fitzpatrick 1984) that the behavior stems slowly walked along the margin of a wide, sand firelane from an ancestralwillingness to perch and forage on borderingdense oak scruband scrubbyflatwoods. The the backsof native mammals, in a low-growing habitat deer stoppedseveral times to browse. During one such that lacks large, diurnal, terrestrial predators. Records stop, a Florida Scrub-Jay(band combination ARZ = exist for Florida Scrub-Jaysperching and apparently 3-year old resident male breeder) flew into view and removing ticks from domestic cattle (Bent 1946) and perched on a pine snag several meters from the deer. feral hogs(Baber and Morris 1980). Dixon ( 1944) and As the deer moved on, the jay made an aborted flight Isenhart and DeSante (1985) observedWestern Scrub- toward it, left the video frame momentarily, then reap- Jays(Aphelocoma californica) doing the same on mule peared and alighted on the back of the deer near its and black-tailed deer (Odocoileushemionus), and the shoulders.The deer showed no indication of surprise, latter authorseven speculatedthat a “cleaning station” but continued to browse, head down. relationship existed between bird and deer. Here we Upon landing the jay immediately picked at the back describean encounterbetween a Florida Scrub-Jayand of the deer as if to take food, then hopped posteriorly an adult white-tailed deer (Odocoileusvirginianus), and to the haunches. There the jay first picked at, then summarize evidence that the behavior is not unusual. sharply pecked at and nibbled, a spot on the deer’s The encounterwas video-taped by a deer hunter while right rear buttock. The sharp peck prompted the deer he sat in a tree overlooking the scene.Our description to lower its haunchesslightly, head still down. The deer remained frozen in this posture for several seconds before picking up its head and casually looking back toward the jay. The jay then hopped forward again to ’ ’ Received 28 August 1995. Accepted 10 January the shoulder region, making a few additional gentle 1996. picks. At this point the deer easily could have brushed 2 Current address: Laboratory of Ornithology, 159 away the jay with its muzzle or antlers, but instead SapsuckerWoods Rd., Ithaca, NY 14859. remained seemingly unperturbed while watching the