136 ShortCommunications andCommentaries [Auk,Vol. 110

The Auk 110(1):136-141, 1993

The Pacific Golden- ( œulva): Discovery of the Speciesand Other Historical Notes

Osc• W. JOHNSON Departmentof Biology,Montana State University, Bozeman, Montana 59717, USA

Knowledgeof a species'discovery and nomencla- shot "4 ploversexactly like our Englishgolden plo- tural chainalong with associatedpeople and events ver" (now Eurasian Golden-Plover, Pluvialisapricaria) adds significantly to our appreciationof it. For most on a small island off the northwestern coast of Java. , thesehistorical details are relatively obscure. In his journal AnimaliumJavanensia, an unpublished Tracingthe pastof the PacificGolden-Plover (Pluvialis work held by the British Museum(Natural History), fulva) toucheson the remarkable enduranceand ac- Solander comparedthe birds to P. apricaria,stating complishmentsof several 18th-centuryornitholo- "agreeswith the descriptionby Brissonand Linnaeus, gists,the perilsof not publishingpromptly, and the exceptbelly is white sprinkledwith a few blackspots work of an early systematistwho becamethe recog- [thesewere birds in prebasicmolt showing remnants nized authorityfor the specieswithout examining of breeding plumage]and perhapsthe spotson the any specimens.This plover has a relatively "distin- tail are morein the shapeof bars."Although Solander guished background" in that it was found and de- seemedaware of differencesbetween these specimens scribed by famous naturalists during Capt. James and P. apricaria,he evidently did not recognize that Cook's explorationsof the Pacific.These extraordi- this plover was a new species.Neither Solander's nary voyagesyielded a wealth of zoologicalinfor- notesnor the firstvoyage specimens (none are known mation,including majorfindings in the areasof to have reachedEngland) have any nomenclaturalor systematicsand distribution. Newly discoveredPa- type status. cificbirds figured prominently in thecyclopedic works The first specimensof taxonomicsignificance were of the time (Latham 1781-1801, Pennant 1785, Gmelin collectedby JohannReinhold Forster,George Forster 1788-1789,Forster 1844); more recently, various dis- (Johann'sson), and AndersSparrman. This illustrious cussionshave emphasizedthe ornithologicalsignif- team of naturalists sailed aboard HMS Resolution dur- icance of Cook's expeditions (Stresemann 1949, 1950, ing Cook'ssecond voyage (1772-1775), an expedition 1975,Lysaght 1959, Wilson 1977, Medway 1979,1981). that involved enormoussweeps of the South Pacific Cook'sthree Pacificvoyages present a fascinatingar- and circumnavigation of Antarctica (for full account, ray of charactersand eventsthat are thoroughlyde- seeBeaglehole 1969). Collection of the type specimen tailed in the monumentalworks of Beaglehole(1962, can be fixed almost to the hour. The bird was collected 1967-1969). From these chronicles and other sources at Matavai Bay, Tahiti on the afternoon of 26 August I haveextracted an historicalperspective of the Pacific 1773 by J. R. Forster.Forster's log indicatesthat the Golden-Plover'sdiscovery, subsequent records and type was shot during the first few hours of 27 August, happenings, and the people involved. I include com- but this was basedon "ship's time," where the day mentswhere the originalfindings can be clarifiedby runs from noon to noon and begins 12 h before the currentknowledge of P. fulvabiology. With respect civil day. In relation to civil time, early on 27 August to geographic distribution of the species,most of is equivalentto the afternoonof 26 August.His jour- Cook'stravels were within the vast winter range of nal containsthe simple notation "shot a new Charad- the Pacific Golden-Plover in the Central and South rius" (Hoare 1982). From this specimen, Forster de- Pacific,and only the third voyagetouched the arctic scribedthe speciesCharadrius glaucopus. Although the breeding grounds.Recent studies of breeding birds descriptionapparently was written at the time of col- in western Alaska(Connors 1983, Connors et al. 1993) lection, the original accountremained unpublished show that P. fulva is a full speciesseparate from the for manyyears. It finally appearedwell after Forster's AmericanGolden-Plover (P. dominica).This separa- death (1798) in DescriptionesAnimalium (1844). tion has been widely accepted(see Johnsonet al. The Resolutiondeparted from Tahiti on 17 Septem- 1989),and is under considerationby the AOU. For ber 1773 and reachedTonga on 2 October. While en morecomplete information on the species(including route, J. R. Forsterrecorded several sightings of plo- , migration, plumages, and behavior), see vers (Hoare 1982).On 26 Septemberhe wrote: "The Connors (1983), Connors et al. (1993), Johnson et al. birds have quite forsaken our Ship. However, one (1981, 1989), Johnsonand Johnson(1983), and John- Landbird, a kind of Sandpiper,such as was seen on son (1985). the 22 & 23d was in the Afternoon hovering about The first specimensever recorded were collected the Ship, & settled twice on it, being quite tired & by the naturalistsaccompanying Cook's first voyage spent:it flew afterwardsaway. It is of the same kind (1768-1771). JosephBanks wrote on 8 October 1770 of Ploveras we observedin Otahaitee(Charadrius glau- (see Beaglehole 1962) that he and Daniel Solander copus)."Based on the chronologyof migration, the January1993] ShortCommunications andCommentaries 137 observedduring this period were probably scribedits legs as "blue" (P. fulva has grayish-black juveniles. Additional plovers were collectedat Tonga legs, but suchcoloration might well be consideredas on 4 October 1773, and at New Caledonia on 6 Sep- a shadeof blue by some observers).Presumably, La- tember 1774 (Hoare 1982). George Forster rendered tham usedthe samereference points when measuring paintings of the speciesat these two sites (Lysaght "vat. A" as he commented that it was "considerably 1959), but there is no known painting of the type less in size" when compared to the type. While it specimenfrom Tahiti. The fate of the type specimen appearsthat "vat. A" was too small to be P. fulva, this and the specimens collected subsequently is un- single measurementwould be inconclusivesince La- known. Most likely they were inadequately pre- tham's specimenmay have been poorly preserved servedand either did not survive the voyage or per- (possiblyit wasshrunken or otherwisedistorted) and ished shortly thereafter. not comparableto Forster'sfreshly collectedbird. Leg DescriptionesAnimalium was essentially complete color, however, is a lesssubjective criterion and "pale when the expedition returned to England in July 1775 yellow" is reasonablygood evidence that "vat. A" (Hoare 1982), but various postvoyagedisagreements was, in fact, not P. fulva. with the British Admiralty prevented its publication Perhaps"vat. A" was the single plover specimen (for discussionsof this troubled period in Forster's from Cook's third voyage,which appearsin the cat- life, see Stresemann 1975, Hoare 1976, 1982). Forster, alog of the Bankscollection (Medway 1979).Although frustratedin his effortsto publish, unfortunately lost records of this bird indicate that it was a Pacific Gold- much of the early recognitionthat was rightfully his. en-Plover, the identification may have been incorrect. Meanwhile, John Latham was writing the GeneralSyn- It appearsthat Banksgave various specimens to Lever opsisof Birds (1781-1801) and incorporated therein (Medway 1979); thus, the bird could have been in most of Forster'sfindings. Thus, Latham published Lever's possessionwhen Latham examined it. Ques- the first account of the Pacific Golden-Plover, which tions concerningthe exactidentity of "vat. A" and he referred to as the "Fulvous Plover" (GeneralSyn- whether it was the same or a different bird than the opsis,vol. 3, 1785). Since the type specimen was un- specimenin Banks'collection are unlikely ever to be available and there was no painting of it, Latham resolved. Almost all of the museum specimensfrom based his description of the type on information Cook's voyages eventually perished (victims of ne- "communicatedby Dr. Forster."As to what was"com- glect and crude procedures),and only a very few are municated," one finds great similarity between La- extant(Sharpe 1906,Medway 1981).Other plover skins tham's descriptionand Forster's(in DescriptionesAn- may have been collected on Cook's third expedition, imalium1844), suggesting that Forsterprovided Latham but only the questionablespecimen mentioned above accessto his original notes. The type specimenwas is known to have survived the voyage.Given such an individual wearing traces of breeding plumage: paucity of specimensand the fact that Latham used its breastand belly were "spottedwith black." Such only "vat. A" to supplementForster's description, it featbering is consistent with the date of collection is entirely possible that Latham never saw an actual and representsan adult bird in prebasicmolt. specimenof P. fulva. Forster'spersonal assistance in Latham'sproject ap- Neither Forster nor Latham emerged as the rec- pearsto have beenonly transitory(Hoare 1976).Much ognized authority for the species.Instead, the dis- of Forster'scollection had by then passedinto the tinction went to Johann Friedrich Gmelin by default. famousprivate museumsof JosephBanks and Ashton Forster lost out becausehis findings were not pub- Lever to which Latham had free access. Banks and lished promptly, Latham becauseof the format he Lever eventually held mostof the skinsand paintings chosefor GeneralSynopsis. The latter was written in from the three Cook expeditions (Sharpe 1906, Ly- English without adherenceto the Linnaean method-- saght 1959,Stresemann 1975, Medway 1979, 1981).It an approach which was unacceptable to the inter- is probable that Latham gained insight as to the fea- national scientific community. Gmelin, then editing turesof P. fulva by viewing one or more of four paint- the 13th edition of SystemaNaturae (1788-1789), in- ings: two by George Forster (mentioned above),and cluded Latham'swork by simply translatingthe ma- two from the third voyage (see below). All of these terial into Linnaean Latin. At that moment, the "Ful- works were then owned by Banks (Lysaght 1959). vousPlover" became Charadrius fulvus. Thus, as others Also, Latham examined a specimen (date and place have noted (Mathews 1931, Stresemann 1949, 1975, of collection unrecorded) in the Leverian Museum, Medway 1981), Gmelin achieved nomenclaturalrec- which he termed "vat. A" of the "Fulvous Plover." ognition for P. fulva and many other bird specieswith However, not all of its features matched the type minimal effort and no direct knowledgeof the spec- specimen:the bird was relatively small ("Length eight imens. inches") and its legs were of the wrong color ("pale Cook's third and last voyage (1776-1780) was pri- yellow"). Forster measuredhis type specimenat 12•A marily for explorationof the North Pacificand Bering inches"from tip of bill to end of claw of middle toe" Sea. The expedition produced an immense array of (this is an accuratedimension basedon personal ex- new information including discoveryof the Hawai- amination of freshly collected specimens),and de- ian Islands (for detailed treatment, see Beaglehole 138 ShortCommunications andCommentaries [Auk,Vol. 110

Fig. 1. GeorgeForster's two paintingsof the PacificGolden-Plover. Upper panel: Bird collected4 October 1773at Tonga;individual nearing completion of prebasicmolt and had only a few flecksof dark breeding plumageremaining on breastand belly. Lower panel: Collected 6 September1774 at New Caledonia;prebasic molt lessadvanced than in other bird and considerablebreeding plumage still in place.Both works have beencropped so as to emphasizethe birds.In theprocess, notations by Forster(faded on originalsand mostly illegiblein my photographs)were removed. For information concerning these notations, see Lysaght (1959). January 1993] ShortCommunications and Commentaries 139

Fig.2. Watercolorsfrom third voyage. Upper panel: Webber's painting of a ploverin basicplumage. Bird couldbe either a juvenileor an adult. After completion ofprebasic I molt and loss of juvenile body feathering (seebelow), age determination isdifficult without the bird in hand.Locality given by artist was "Sandwich Islands,1777"; however, this is incorrectsince the expedition did not sight Hawaii until 18 January 1778. Lowerpanel: Ellis' rendering of a ploverfrom Christmas Island. Based on voyage's itinerary, bird collected between25 December1777 and 2 January1778. It appearsthat specimen had barred flanks, which would indicatejuvenile plumage. Although collection period is somewhat late for such feathering, it is possible that thebird was a youngindividual in itsfirst wintering season. For additional information on these paintings, see Lysaght(1959) and l•edway (1981). 140 ShortCommunications andCommentaries [Auk,Vol. 110

1967). Although there were no official naturalists In addition to the various third-voyage sightings aboard, extensivecollections, drawings, and records described above, P. fulva was further documented of birds were obtained by several individuals pos- through the artistry of William Ellis and JohnWebber sessingscientific skills. Particularly notable were Wil- (a landscapeartist aboard the Resolution).Each pro- liam Anderson (ship's surgeon)and William Ellis (as- duced a single watercolor of the bird. The Ellis work sistantsurgeon). Anderson demonstratedremarkable and the two paintings by Forster (from the second abilities in ornithology (Iredale 1938, Stresemann voyage) are in the collectionsof the British Museum 1950).Regrettably, he died during the voyageand his (Natural History), Webber's painting is held by the losswas "very much to the disadvantageof zoological British Museum (Lysaght 1959). All four works are success"(Stresemann 1949). shown in Figures1 and 2. This is the first time they Within the journalsfrom the third expedition(most have been reproduced. have been published and/or examined in detail; see I am grateful to David G. Medway who generously Ellis 1782,Beaglehole 1967, Medway 1979,1981)there sharedhis extensiveknowledge of ornithologyin the are a number of references to plovers. While these Cook era and supplied copied material from the un- sightingsseem reliable, confusionwith other species publishedSolander manuscript. Alan Knox provided of shorebirdscannot be ruled out in every instance. early guidance in locating the paintings by Forster, JamesCook recorded plovers at Sledge Island near Ellis, and Webber. Officials at the British Museum present day Nome, Alaska on 5 August 1778, and saw (Natural History) and the Trusteesof the British Mu- flights of "birds like Plover" (probably adult P. fulva seum kindly allowed these materials to be repro- in migration) from mid- to late August 1778 on the duced. Peter G. Connorsand C. Stuart Houston pro- Bering Sea. During this period, Cook ponderedthe vided helpful commentson the manuscript. This southwardmovements of birdsand speculated:"Does investigationwas donein associationwith other stud- not this indicate that there must be land to the North ies of the PacificGolden-Plover funded mostlyby the where thesebirds retired in the proper seasonto breed National GeographicSociety. Additional support was and were now returning to a warmer climate." Cook's provided by the Hawaii Audubon Societyand Moor- insight is perhapsthe first recordedstatement con- head State University (Minnesota) through a faculty cerningmigration in the northern Pacificregion. There researchgrant program. is a notable discrepancyas to the number of plovers observed at Sledge Island. JamesKing (one of the LITERATURE CITED expedition'sofficers) wrote in his journal that he and Cook had walked together over the area. Whereas ALCORN,R. 1988. AWSG regular countspro- Cook noted "a few Plover," King recorded"flocks of ject interim report to June 1987:Migratory wad- Plovers." William Anderson observed birds which ers. Stilt 12:7-23. "seem'd to be of the Plover kind" on 20 March 1777 BARTER,M.A. 1988. Biometrics and moult of Lesser between New Zealand and the Cook Islands (these Golden PloversPluvialis dominica fulva in Victoria. may have been spring migrants;northward move- Stilt 13:15-19. ments from the far Southern Hemisphere start rela- BEAGLEHOLE,J. C. (Ed.) 1962. The Endeavourjournal tively early in the year; see Lane 1987, Alcorn 1988, of JosephBanks 1768-1771, 2 vols. Angus and Barter 1988),noted plovers at PalmerstonAtoll on 17 Robertson,Sydney. April 1777,and describeda plover "spottedwith yel- BEAGLEHOLE,J. C. (Ed.) 1967-1969. The journalsof low" at Tonga on 11 July 1777 (most likely an over- Captain JamesCook on his voyagesof discovery. summering individual). William Ellis recorded plo- Vol. 1 (1968), The voyage of the Endeavour1768- vers on ChristmasIsland in late December to early 1771;vol. 2 (1969), The voyage of the Resolution January 1778,at Nootka Sound (on Vancouver Island) and Adventure1772-1775; vol. 3 (1967),The voy- in late April 1778 (the location might indicate birds age of the Resolutionand Discovery1776-1780. from California wintering groundsrather than trans- Hakluyt Society, Cambridge Univ. Press, Cam- oceanicmigrants), on UnalaskaIsland in late October bridge. 1778 (migrating juveniles), and on the island of Ha- CONNORS,P.G. 1983. Taxonomy, distribution, and waii in the early months of 1779. During the 1779 evolutionof goldenplovers (Pluvialis dominica and stay in Hawaii, Charles Clerke (who assumedcom- Pluvialisfulva). Auk 100:607-620. mand of the Resolutionfollowing Cook's death on 14 CONNORS, P. G., B. J. MCCAFFERY,AND J. L. MARON. February 1779) also noted the occurrenceof plovers. 1993. Speciation in golden-plovers, Pluvialis Latham's(1785) remark that "our last voyagers"ob- dominicaand P. fulva:Evidence from the breeding servedplovers "at Owhyhee" wasevidently basedon grounds. Auk 110:9-20. Ellis' 1779records (Medway 1981).Since the context ELLIS,W. 1782. An authentic narrative of a voyage of Latham'scomment had to do with golden-plovers performed by Captain Cook and Captain Clerke in general, he apparently was unaware that his "Ful- in His Majesty'sShips Resolutionand Discovery vous Plover" from Tahiti and the plover in Hawaii during the years 1776,1777, 1778,1779, and 1780. were of the samespecies. 2 vols. Robinson, Sewall, and Debrett, London. January1993] ShortCommunications andCommentaries 141

FORSTER,J.R. 1844. DescriptionesAnimallure quae 3 vols.with 2 supplements.Leigh and Sotheby, in itinere ad marls australistetras per annos 1772, London. 1773 et 1774 susceptocollegit observavitet de- LYSAGHT,A. 1959. Some eighteenth century bird lineavit JoannesReinoldus Forster (H. Lichten- paintingsin the library of Sir JosephBanks (1743- stein, Ed.). AcademiaLitterarum Regia, Berlin. 1820). Bull. Br. Mus. Nat. Hist. (Historical Ser.) GMELIN,J.F. 1788-1789. SystemaNaturae, 13th ed. 1:251-371. Beer,Lipsiae. MATHEWS,G.M. 1931. John Latham (1740-1837): An HOARE,M.E. 1976. The tactlessphilosopher Johann early English ornithologist.Ibis 1 (13th ser.):466- Reinhold Forster (1729-98). Hawthorn Press, 475. Melbourne. MEDW^Y,D.G. 1979. Someornithological results of HO^RE,M. E. (Ed.) 1982. The Resolutionjournal of Cook's third voyage. J. Soc. Bibliog. Nat. Hist. JohannReinhold Forster 1772-1775. Hakluyt So- 3:315-351. ciety, Cambridge Univ. Press,Cambridge. MEDW^Y, D. G. 1981. The contribution of Cook's IREDALE,T. 1938. WilliamAnderson--Ornithologist. third voyageto the ornithologyof the Hawaiian Islands. Pac. Sci. 35:105-175. Emu 38:60-62. JOHNSON,O. W. 1985. Timing of primary molt in PENNANT,T. 1785. Arctic zoology. Vol. 2, Birds. first-yeargolden-plovers and someevolutionary Hughs, London (reprinted by Arno Press,1974). implications.Wilson Bull. 97:237-239. SHARPE,R. B. 1906. Birds. Pages79-515 in The his- tory of the collectionscontained in the natural JOHNSON,O. W., ^ND P.M. JOHNSON.1983. Plumage- history departmentsof the British Museum, vol. molt-agerelationships in "over-summering"and 2. Trustees of the British Museum, Longmarts, migratoryLesser Golden-Plovers. Condor 85:406- London. 419. STRESEMANN,E. 1949. Birds collected in the North JOHNSON,O. W., P.M. JOHNSON,AND P. L. BRUNER. Pacificarea during Capt. JamesCook's last voyage 1981. Wintering behavior and site-faithfulness (1778 and 1779). Ibis 91:244-255. of golden plovers on Oahu. Elepaio 41:123-130. STRESEMANN,E. 1950. Birds collectedduring Capt. JOHNSON,O. W., M. L. MORTON, P. L. BRUNER,AND P. JamesCook's last expedition (1776-1780). Auk 67: M. JOHNSON.1989. Fat cyclicity, predicted mi- 66-88. gratory flight ranges, and features of wintering STRESEMANN,E. 1975. Ornithology from Aristotle to behavior in Pacific Golden-Plovers. Condor 91: present. Harvard Univ. Press,Cambridge. 156-177. WInSON, E. 1977. Observations of Hawaiian avifauna LANE, B. A. 1987. Shorebirds in Australia. Nelson, during Cook'sexpeditions. Elepaio 38:13-18. Melbourne. LATHAM,J. 1781-1801. A generalsynopsis of birds. Received2 October1991, accepted 22 February1992.

The Auk 110(1):141-145, 1993

Comparison of Rod/Cone Ratio in Three Speciesof Shorebirds Having Different Nocturnal Foraging Strategies

Luz MARINA ROJASDE AZUAJE,•'3 SUSAN T•a,' AND RAYMOND MCNEIL 2 •Institutode Investigacionesen Biomedicinay Ciencias Aplicadas, Universidad de ¸riente, Cumand, Sucre, Venezuela; and 2Dipartementde sciencesbiologiques, Universit[ de Montrial, C. P. 6128, Succ."A", Montreal, QuebecH3C3J7, Canada

Nocturnality, the habit of being activeduring dark- see McNeil 1991, McNeil et al. 1992, 1993). Two basic ness,is a characteristicof some aquatic birds, partic- types of foraging techniquesare used by shorebirds: ularly those in the shorebird families (McNeil and visual searchingfor prey or indicationsof their pres- Robert 1988, Robert and McNeil 1989, Robert et al. ence, on or near the surface (e.g. Pluvialisand Cha- 1989; for detailed review concerning aquatic birds, radriusplovers); and probing with the bill for buried prey that are detectedby tactile and taste cues(e.g. Short-billed Dowitcher, Limnodromusgriseus). Some 3 Present address:D6partement de sciencesbiolo- speciesfeed exclusivelywith the sameforaging tech- giques,Universit• de Montr6al, C. P. 6128,Succ. "A", nique both by night and by day (e.g.visual searching Montr6al, Quebec H3C 3J7, Canada. by Pluvialisand Charadriusplovers, or tactile probing