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The many unknowns about and of the world: introduction to a wealth of research opportunities highly relevant for shorebird conservation

TheunisPiersma, Popko Wiersma& Jan van Gils

Piersma, T., Wiersma, P. & van Gils, J. 1997. The many unknownsabout plovers and sandpipersof the world: introductionto a wealth of research opportunitieshighly relevant for shorebird conservation. Study Group Bull. 82: 23-33. We reviewthe current information,especially with respectto conservationbiology, about the () and (Scolopacidae) families, based on informationassembled for the -chaptersin the Handbook of the of the World, Volume 3 (del Hoyo et al. 1996). The existing information about 155 is summarized in a comprehensivetable (Appendix1), and some informativestatistics have been derived. Existingknowledge is very unequallydistributed. For example, for those plovers and sandpipersbreeding in only, nothingis known about the demographyof of the species. Species confinedto and are slightlybetter known, but also here demographicand feeding ecological knowledgeis absent in most cases. Best studied are species in the remaining regions of the world, but even for species breeding in any demographicalknowledge, so criticalfor soundconservation practise and management,is lackingfor a third of the species. There are no fewer than 27 plover and sandpiper species about which virtually nothing is known. Inland species such as several , plovers, and with restricted distributionin South America, Africa and Asia, feature prominentlyin this list; a third of these species are known to face conservationproblems. There is work to be done by waderologists all over the world.

T. Pierstoa & J. van Gils, Netherlands Institute for Sea Research (NIOZ), P.O. Box 59, 1790 AB Den Burg, Texel, The Netherlands;E-mail: theunis•nioz. nL P. Wiersma,Zoological Laboratory, University of Groningen,P.O. Box 14, 9750 AA Haren, The Netherlands.

INTRODUCTION hope that the knowledge eventually gained will help to safeguard endangered wader species in due course. Duringthe preparationof the speciesdescriptions, species distributionmaps and family texts of the two most diverse families of (the Charadriidaeand the Scolopac- METHODS idae) for the Handbookof the Birdsof the World,Volume 3 (del Hoyo, Elliott& Sargital 1996), the three of us For the preparationof the species-texts(Piersma & assembled a lot of information in a fairly comprehensive Wiersma 1996; Piersma et al. 1996), most of the regional and systematicway (Piersma & Wiersma 1996, Piersma handbookswere examined, all "Recent publicationson et al. 1996). Ouringthis process,we became acutely waders"published in the Wader Study Group Bulletinwere aware of the paucity of informationon many species, scanned, and several computerised literature searches suggestingmuch scopefor basic investigationsin most were made. The relevant information was assembled partsof the world, and we also realizedthe need for under the headings "","Distribution", further integrativeand comparativestudies on the biology "Descriptivenotes", "", "Food and Feeding", of these families. In this brief review we present a "Breeding","Movements", "Status and Conservation"and tabulated summary of our findings(Appendix 1). As the "Bibliography".For the compilationin Appendix 1 we used knowledgebase for the otherwader families may not be our original, unedited bibliographiesand counted the directlycomparable, we have refrainedfrom the number of publicationsspecifically devoted to the biology temptationto do a similar exercisefor them as well. of the species concernedto assign it to one of the six categoriesof the publicationrecord. To indicatethe strengthand weaknessesin the worldwide knowledgeabout ploversand sandpipers,we have On the basis of the distributionmap and the "distribution" summarizedthe informationin Appendix1 furtherin a sectionwe assigned each species to one or more series of tables. Of most relevance, of course, is an continents,adding Oceania (the islands of the Pacific identificationof the state of knowledgeabout rare and Ocean) as the seventh region. In AppendixI we then endangeredwader species. If this reviewstimulates listed the official conservationstatus accordingto the relevantexplorations and focussessome of the necessary IUCN criteria as adopted and assigned by BirdLife field and desk efforts, we have achieved our aims. We International(Collar eta/. 1994). Many speciesgive no

23 immediate conservation concern and are called "Not- 3) reproductivebiology (mating system, breeding GloballyThreatened". Species that may have a problem biology,parent-offspring relationships), are "Near-Threatened". Accordingto the IUCN criteria, 4) migrationsystem (use of flyways by different speciesin dire straightsare either"Vulnerable"(awarded a segments of population,geography of wintering and 10% chance of going extinctin the next 100 years), breedinggrounds, (re-) fueling strategies), and "Endangered"(with a 20% chanceof goingextinct in the next20 years) or "Critical"(with a 50% chanceof going 5) food and foraging. extinctwithin five years). Obviously,there is a degree of subjectivityin making these assignments,but this seems unavoidable. The Apartfrom the officialconservation status, Appendix 1 patternsof No's and Yes's shouldnevertheless give a fair givesa listingof the degreeof knowledgeabout five topics pictureof the state of our knowledgebase. that are most relevant for conservationbiology. For each specieswe haveasked ourselves, on the basisof the information assembled for the Handbook of the Birds of the World, whether nothing("No"), something ("+") or quite TAXONOMIC DIVERSITY AMONG PLOVERS AND much ("Yes")is knownabout respectively SANDPIPERS 1) populationsize, The diversityamong the Charadriidaeis illustratedin 2) demographicstructure (age composition,sex ratio, Figure 1. recruitment,survival and mortalityfactors),

MagellanicPlover 1 species (Pluvianellus) lapwings 25 species (Vane#us,Erythrogonys)

plovers 41 species (Pluvialis,Charaddus, Elseyornis, Peltohyas,Anarhynchus, Phegornis,Oreopholus)

Figure1. Subdivisionofthe plovers, Charadriidae, inthree subfamilies, illustrating thediversity ofthis wader family, with listings of the currently recognizedgenera, and total numbers of extantspecies. The white-on-black name is that of the family, whilst the shaded boxes give sub-family names.This is a black-and-whitereprint (by permission ofthe editors) ofa figureby ,•,ngels Jutglar & Etel Vilar6 in del Hoyo et al. (1996: p. 384).

With the exceptionof the MagellanicPlover (Pluvianellus outgroupto the othertwo, havingderived from a common socialis;subfamily Pluvianellinae), clearly not belongingto ancestorearly on. Perhaps,Pluvialis should not be the cladeof ploversand usuallyawarded family status, includedin the subfamilyCharadriinae, but rather have a biochemicalevidence suggests that all modernplovers subfamilyof its own. It has beensuggested several times share the same commdn ancestor. Althoughthe family is that the ploversare a familythat originatedat low commonlydivided into two subfamilies,the lapwings latitudesin the southernhemisphere, the regionwhere (Vanellinae)and the true plovers(Charadriinae), the Grey mostspecies are aroundtoday. The ploverfamily should andgolden plovers of the genusPluvialis may be an have evolved under rather arid, semidesert conditions,

24 specializingon smallprey that are mostactive at night. single subfamily of -alikes,the exact positionof the Onlythe genusPluvialis, of whichall four speciesbreed in three (Limnodromus) remains unresolved. tundra areas aroundthe Ocean, may have its origin Most previousauthors have listedthem with the snipes, in the northern hemisphere. but others have put them with the tringinesandpipers, and for both hypothesesthere is somethingto say. The The greatdiversity of the sandpiperfamily, Scolopacidae, tringine-wadersare very likelyto be composedof different which consists of some six subfamilies (woodcocks, evolutionarylineages. Recent work suggeststhat the snipes,, sandpipers, tringine-waders and shanks and tattlers may be closely related to the ),with the Tringinaebeing subdivided further in phalaropes,and that this combined lineage is a sister & ,shanks and Polynesiansandpipers, is group to the woodcocksand snipes. The godwits illustratedin Figure 2. There is little doubtthat the (Lirnosa)and the curlews(Nurnenius and Bartrarnia) Scolopacidaeis monophyletic,but there are problemswith appear to be two independenttaxa, quite unrelatedto the the biologicalsubdivision of this variablefamily. Four of rest of the Tringinae, possibly branching off at the base of the subfamilies(the woodcocks,turnstones, sandpipers, the scolopacidfamily. The positionof the Tuamotu and phalaropes)are almostcertainly monophyletic. Apart Sandpiperand its two extinctrelatives remains unresolved from a possiblemerger of woodcockswith snipesinto a in view of a lack of modern phylogeneticstudies.

i•.••, .....:•:. •..-'':•',,•'• -.-.-.• - ;::•,•,•....•, .,,•r • .•-•.•. - .

woodcocks snipesand dowitchers turnstones sandpipers phalaropes 6 species 21 speci• 2 species 24 spedes 3 species (Scolop•) (C•n•o•ha, Lymnoc•tes, (Arenaria) (Aphriza,, (Steganopus,Phalaropus) ,Limnodrom•) Eurynorhynchus, Lirnicola,Micropalarna, Tryngites,Philornachus)

I NUMENIINIII

•••:•"'• .•... ;?•...... godwitsand curlews shanks Polynesiansandpipers 13 species 16 species 1 species (Lirnosa,Nurnenius, (,Xenus, , (Prosobonia) Bartrarnia) Heteroscelus,Catoptrophorus)

Figure2. Subdivisionofthe sandpiper-family, Scolopacidae, insix subfamilies (and the subfamily Tringinae inthree tribes), illustrating thegreat diversityofthis wader family, with listings ofthe currently recognized genera, and total numbers ofextant species. The white-on-black name is thatof the family, whilst the shaded boxes give sub-family names, and open boxes give names of tribes. This is a black-and-whitereprint (by permissionofthe editors) ofa figureby Francesc Jutglar and Etel Vilar•) in del Hoyo et al. (1996: p. 445).

25 OVERVIEW OF PUBLISHED INFORMATION

The four best known plover and sandpiper species in the Table 1. The four best knownplover and sandpiperspecies of the world are the , the Black-tailedGodwit, the Red world. These speciesscored a "Yes"under each of the five Knot and the (Table 1), each of which scored a categoriesof knowledgelisted in Appendix1. "Yes"for all five knowledge-categoriesin Appendix 1. English name Latin name Breeding area Even thoughvery well-studied,our knowledgeabout these Wrybill Anarhynchus NewZealand speciesis, not even remotely, approachingthe degreeof frontalis understandingof the biologyof Eurasian Oystercatcher (Haematopusostralegus, Haematopodidae), about which Black-tailed Limosalimosa temperateEurope & well over 500 papers have been published(Goss-Custard Asia 1996). The five most abundantlyresearched plover and Calidtfscanutus circumpolar sandpiperspecies in the world are NorthernLapwing, , Red Knot, Dunlin and (Table 2). Dunlin Calidtfsalpina circumpolar Even for those species,large parts of the biologyare not very well known,often becausethe particularsubject is difficultto study in the particularspecies (e.g. food and feedingin ). Notethat in the case of the two Table 2. The five mostabundantly researched plover and sandpiper Coenocorypha-snipesfrom the oceanic islandsof New speciesof the world. These speciesscored an "F"for their Zealand, a publicationrecord of "A"is not indicativeof a publicationrecord: more than 50 topical(i.e. devoted)publications by poorlystudied species. In fact, these two specieshave 1995. had detailed attention, and the results were publishedin a English name Latin name Breeding area few excellentand detailedpapers (e.g. Miskelly1990). Northern Vanellusvanellus temperateEurope & Asia Goingfrom the well-researchedto the poorlyresearched side of the ploverand sandpiperrealm, Table 3 liststhe Common Tt•ngatotanus temperateEurope & degreeof lack of knowledgeabout different topics for Redshank Asia speciesbreeding in differentparts of the world. (Note that Red Knot Caliddscanutus circumpolar the specieshave been arrangedhierarchically with respect to best studiedregion [Europe], and that the geographical Dunlin Caliddsalpina circumpolar categoriesin Table 3 are not mutuallyexclusive.) Overall, Ruff Philomachuspugnax temperateEurope & populationsize is completelyunknown in about half the Asia species. Knowledgeabout the demographyof wader speciesscores even more poorly,with 70% (!) of the world's plover and sandpiperspecies not being covered. The remainingthree knowledge-categoriesfare better, with only a third to a quarterof the speciescompletely lackinginformation.

Table3. Relativelack of knowledgeabout plovers and sandpipers of theworld, broken down by geographical (breeding) region. For each of the fivecategories of knowledge,the percentageof speciesthat scored "No's" according to informationinthe Handbook of theBirds of the Wodd, Vol.3 (Appendix1) is given.Species are ordered according to breedingorigin, with the least studied regions being followed by the better studied regions.Species that breed uniquely in SouthAmerica, Africa, Asia and + Oceaniaare examined first. Of the remainingspecies, those thatbreed in NorthAmerica (but not in Europe,including Greenland) are examined next, finally followed by those that (also) breed in Europe. Breedingarea No Population Demography Reproduction Migration Food& Overall species size foraging score Overall 155 46 % 70 % 23 % 22 % 32 % 39 %

South America 15 87 % 100 % 73 % 53 % 87 % 80 %

Africa 19 95 % 89 % 21% 37 % 58 % 60 %

Asia 35 51% 86 % 34 % 31% 49 % 50 %

Australia& Oceania 16 0 % 63 % 19 % 25 % 25 % 26 % NorthAmerica 34 38 % 71% 9 % 0 % 3 % 24 % Europe 36 28 % 33 % 0 % 0 % 8 % 14%

The existingknowledge is very unequallydistributed any of the species. Speciesconfined to Africaand Asia (Table3). Of the ploversand sandpipersbreeding in are slightlybetter known, but also heredemographic and SouthAmerica only, 80% of the possibleslots scored a feedingecological knowledge is absentin most cases. "No",with nothing being known about the demographyof Best studiedare speciesin the remainingregions of the

26 world, but even for species breedingin Europe any PLOVER AND SANDPIPER SPECIES OF demographicalknowledge, so criticalfor sound IMMEDIATE CONSERVATION CONCERN conservationpractise and management, is lacking for a third of the species. It is noteworthythat the Australians Of the 155 speciesin Appendix1, 33 face the danger of are doing very well with respect to population passingaway and three have gone extinctalready assessments, although again little is known about (Table 5). In this paragraphwe Idiscussthe species that demographic aspects of the majority of their species. are of most concern, and therefore need the particular attentionof conservationistsand waderologists. There are no fewer than 27 plover and sandpiper species about which virtually nothingis known (Table 4). Inland The one recognizedplover species that certainlywill not speciessuch as several lapwings,plovers, woodcocks make it into the 21th century is the Javanese Wattled and snipeswith restricteddistribution in South America, Lapwing, historicallyan endemic of the Indonesian Africa and Asia, feature prominentlyin this list. The Archipelago. It was last observedon the southeastcoast remoteness of many of the regions where the species of Java in 1939. JavaneseWattled Lapwingsprobably occur is obviouslya factor that has inhibitedthe study of succumbedunder the combinedpressures of degradation their ecology. Nevertheless,it is opportunefor work to be of breedinghabitat by intensifiedagricultural practices and done on these species, since a third (nine out of 27) of the hunting. Other island populationsof plovers are birds listed as unstudied are known to face conservation threatened with extinctiontoo. The two populationsof problems.

Table4. Hitlistof the 27 extantplover and sandpiperspecies about which virtually nothing is known. This selectionis basedon data in Appendix 1, and givesspecies for which the sum of the scoresfor the fivedifferent kinds of knowledge(where No = 0, _+= 1 and Yes = 2) is zero or one. Speciesprinted in bold belongto one of the categoriesof (near)threatened species. English name Latin name Breeding area

Spot-breastedLapwing Vane#usmelanocephalus Africa NotGlobally Threatened Brown-chestedLapwing Vane#ussuperciliosus Africa NotGlobally Threatened PiedLapwing Vane#uscayanus SouthAmerica NotGlobally Threatened AndeanLapwing Vane#usresplendens SouthAmerica Not GloballyThreatened Long-billedPlover Charaddusplacidus Asia NotGlobally Threatened Black-banded Plover thoracicus Africa Vulnerable Three-bandedPlover Charad#ust#colla#s Africa Not GloballyThreatened JavanPlover Charad#usjavanicus Asia Not GloballyThreatened MalaysianPlover Charad#usperon# Asia NotGlobally Threatened Chestnut-bandedPlover Charad#uspallidus Africa Not GloballyThreatened Puna Plover Charad#usalticola SouthAmerica Not GloballyThreatened Diademed Plover Phegornis mitchelli South America Near Threatened Tawny-throatedDotterel Oreopholusru#collis SouthAmerica Not GloballyThreatened Amami Scolopax mira Asia Vulnerable RufousWoodcock $colopaxsaturata Asia Not GloballyThreatened SulawesiWoodcock Scolopax celebensis Asia Near Threatened MoluccanWoodcock Scolopaxrochussenii Asia Vulnerable SolitarySnipe Gallinagosolitaria Asia Vulnerable Gallinagonemoricola Asia Vulnerable AfricanSnipe Gallinagonigripennis Africa NotGlobally Threatened MadagascarSnipe Gallinagomacrodactyla Africa NotGlobally Threatened SouthAmerican Snipe Gallinagoparaguaiae SouthAmerica NotGlobally Threatened NobleSnipe Gallinagonobilis SouthAmerica NotGlobally Threatened GiantSnipe Gallinagoundulata SouthAmerica NotGlobally Threatened FuegianSnipe Gallinagostricklandii South America Near Threatened AndeanSnipe Gallinagojamesoni SouthAmerica NotGlobally Threatened Imperialsnipe Gallina•1oimperialis SouthAmerica NearThreatened

Red-breastedPlover, living on Stewart Island (65 birds) St. Helena Plover, of which about 300 were alive during and North Island (1400 birds) of , have the last survey in 1993. The Black-bandedPlover, an shown drastic declinesin numbers and in range over the endemic of Madagascar, is consideredvulnerable. The past 150 years. As a resultof predationby introduced speciesis limitedto the dry and saline lowlandsin and cats, the effectivebreeding population on Stewart southwesternMadagascar, but may not be as rare or as Island is reducedto 12 pairs. AnotherNew Zealand endangeredas previouslysuggested (F. Hawkins,VVVVF- endemic, the Shore Plover, lives on a small (2 km2) island Madagascar,pers. comm. December1996). in the Chatham group. It is doing slightlybetter with 40- 50 breedingpairs. The populationhas been stable Althoughplover populationsliving on islands may run the between1969 and 1993 and is probablyconstrained by greatest risk of extinctiondue to their restricted the availabilityof suitablehabitat ever sinceShore Plovers distribution,there are several continental and more have become extinct on North and more than widespreadplover species that give cause for concern. 100 years ago. Habitatalso seemsthe limitingfactor for Sociable Lapwingswere once quite widespreadon the the third ploverspecies that is regardedendangered, the steppesin southernRussia and Kazachstan. They are

27 now in sharp declinedue to changes in land use for mightwell be in the dangerzone. One is the Grey-headed agriculturalreasons, and, perhaps,desertification. Lapwingof Manchuriaand neighbouringareas of SociableLapwings are consideredvulnerable. On the and far-eastern Russia. There are indications that the North American subcontinenttwo vulnerable plover populationof less than 20 000 birds is decreasing. Land- speciesare the PipingPlover, a of open, oftensaline, use changes and the applicationof pesticidesand lowland , and the Mountain Plover, a bird breeding herbicideson the tropical southeast Asian wintering and winteringin shortgrassprairies. In the last century grounds,could form a seriousthreat for this species.' The both specieswere still very widespreadbut are now in Long-billedPlover has a slightlylarger but overlapping steep declineas a consequenceof changesin land use range, perhaps an even smaller world population(less and, perhaps, hunting. The two remainingvulnerable than 10 000 birds), and very similar problems. In both ploversare the HoodedPlover, a southAustralian species,the resident populationsin Japan are of small endemic shorelinespecies that is threatened by an and unknown size, but may be relatively safe. Grey- increased human use of beaches and nest predation by headed Lapwingsand Long-billedPlovers require urgent and introducedmammals, and the Wrybill from New attention,and so does one more residentspecies from Zealand. Hunted until the 1940s for sport, the population southeastAsia, the coastally living Malaysian Plover. Its of Wrybillsexpanded until the 1960s, when it stabilized total populationsize is also smaller than 10 000 birds, but between 5000 and 6000 birds. However, the entire only410 individualswere countedduring the most recent populationmay be at risk if the breedingsites on braided international Asian waterfowl census. rivers in the Canterburyand Mackenzie Basins of South Island were to be modified by the developmentof In South America the Diademed Plover, an endemic hydroelectricschemes. restrictedto the Andes, has an unknownbut probably small populationsize. It lives in areas that are difficultto There are a few widespread inland east Asian plover access, and its status needs clarification. The endemic speciesabout which preciouslylittle is known, but which

Table 5. The 36 extinct,critical, endangered, vulnerable and (near) threatenedspecies of ploverand sandpiperof the world.

English name Latin name Breeding area Conservation status

JavaneseWattled Lapwing Vanellusmacropterus Asia Extinct White-wingedSandpiper Prosobonialeucoptera Oceania Extinct MooreaSandpiper Prosoboniaellisi Oceania Extinct

Eskimo Numenius borealis Critical Slender-billed Curlew Numenius tenuirostris Asia Critical

Red-breastedPlover Charadriusobscurus Oceania Endangered St Helena Plover Charadriussanctaehelenae Africa Endangered Shore Plover Charadriusnovaeseelandiae Oceania Endangered Nordmann'sGreenshank Tringaguttiler Asia Endangered TuamotuSandpiper Prosoboniacancellata Oceania Endangered

SociableLapwing Vanellusgregarius Europe,Asia Vulnerable PipingPlover Charadriusmelodus NorthAmerica Vulnerable Black-banded Plover Charaddus thoracicus Africa Vulnerable Mountain Plover Charaddus montanus North America Vulnerable Hooded Plover Charadrius rub#co#is Australia Vulnerable Wrybill Anarhynchusfrontalis Oceania Vulnerable AmamiWoodcock Scolopaxmira Asia Vulnerable MoluccanWoodcock Scolopaxrochussenii Asia Vulnerable ChathamSnipe Coenocoryphapusilia Oceania Vulnerable SolitarySnipe Gallinagosolitaria Asia Vulnerable WoodSnipe Gallinagonemoricola Asia Vulnerable Bristle-thighedCurlew Numeniustahitiensis NorthAmerica Vulnerable Spoon-billedSandpiper Eurynorhynchuspygmaeus Asia Vulnerable Broad-billedSandpiper Limicolafalcinellus Europe,Asia Vulnerable

DiademedPlover Phegomismitchelli SouthAmerica N ear Threatened MagellanicPlover Pluvianellussocialis SouthAmerica Near Threatened SulawesiWoodcock Scolopaxcelebensis Asia Near Threatened SubantarcticSnipe Coenocoryphaaucklandica Oceania Near Threatened Latham'sSnipe Gallinagohardwickiii Asia Near Threatened GreatSnipe Gallinagomedia Europe Near Threatened FuegianSnipe Gallinagostricklandii SouthAmerica Near Threatened ImperialSnipe Gallinagoimperialis SouthAmerica N ear Threatened Asian Umnodromussemipalmatus Asia N ear Threatened HudsonJanGodwit Limosa haemastica North America Near Threatened FarEastern Curlew Numeniusmada•lascariensis Asia Near Threatened

28 "plover"from Patagonia, the Magellanic Plover, may have The categoryvulnerable counts two long-distance sufferedbadly from the introductionof sheep in Tierra del migratingsandpipers, the Spoon-billedand Broad-billed Fuego 100 years ago. MagellanicPlovers make no Sandpipers,the Bristle-thighedCurlew breeding in attemptsto distract potentialpredators or herbivorous and winteringon islandsspread out over the Pacific, and intruders from their nests or chicks. Since the shores of four speciesof sedentarysnipes and woodcocks. The pondswhere they breed are paths for hordes of sheep, populationsof vulnerablemigrant sandpipersand curlews trampling is still an importar)tcause of loss. With a all numberseveral to many 1000 individuals. The total populationof less than 1500 individuals,it is is restricted to two islands in the potentiallya vulnerable species. Certain management Mollucas,Indonesia. Only knownfrom eight specimens, practices,for instancekeeping sheep away from breeding the most recentof which was collectedin 1980, they may sites during the nesting season, may both be practical, possiblyhave been observed recently on one of the and critical to safeguard this weird and wonderful islands. The , restricted to broadleaf shorebird species. forest on several small islands in southernJapan, is probablyless rare and may count several 1000 birds. It is Althoughthe status of no fewer than 21, or almost a neverthelessseriously threatened by deforestation and quarter, of the living Scolopacidaegives reasons for predationby newly released mongooses. A thousand concern,there are only two knownextinctions in the past pairs of are confinedto two small few centuries. Both are members of the tringine predator-freeislands in the group, New Prosobonia,and they were lumped as a single species Zealand. The Wood Snipe is a bird of alpine meadows until 1991. The two species of White-winged Sandpiper, with scattered scrubs and streams. With an unknown P. leucopteraand P. ellisi,lived on the adjacent islandsof populationsize, it is confinedto the Himalayan region, Tahiti and Moorea in the Society Archipelago, escaping the harsh winters by a short southward respectively. Based on a few skinsand several drawings, migration to the lowlands of south and southeast Asia. the two sandpipers were differentwith respect to several Another snipe of high slopes close to the timberline, the plumage characteristicsand in the shape of the bill. If , was thought to be extinct for over 100 they indeed lived as residentsalong mountainstreams, it years until it was rediscoveredin 1967 in the Andean is quite likelythat the two populationswere geographically highlandsof , about 2000 km south of the locality isolated in spite of their proximity. The sandpipersof from which it was originally known. Tahiti and Moorea passed away unnoticedin the late 1800s, perhaps as a consequenceof the introductionof goats, pigs and rats on the islands. A close relative, the NEED FOR COMPARATIVE AND INTEGRATIVE Tuamotu Sandpiperis still distributedwidely but thinly STUDIES over a 3 700 km piece of ocean within the Tuamotu Archipelago,east of Tahiti in FrenchPolynesia. It is With such a diverse group of species,the Charadriidae consideredendangered. Several hundred individualslive and Scolopacidaenot only offer examples of conservation scattered over two handfuls of different islands, all of problems, both families and the knowledgeassembled which are isolated and rarely visited, and none of which about the specieswithin the families offer much scope for has rats. concerns other than . The opportunitiesfor good studiesin comparativebiology A similar status has been awarded to the migratory offered by shorebirdshave been recognizedwidely (see, Nordmann's Greenshank, breeding at the edge of for example, Myers 1981; Reynolds& Szbkely 1997; peatmoss/larchbog forestsnear coastal saltmarshes Szbkely & Reynolds 1995), but there undoubtedlyremains around the Sea of ¸khotsk in far-eastern Russia and much more to de done. With respectto comparative winteringin coastal sites in southAsia. With a world demography of plovers and sandpipers, embarrasingly populationof lessthan 1000 birds,Nordmann's little has been achievedsince Hugh Boyd'sfirst review in Greenshankis threatenedby habitat loss on breedingand 1962. The wealth of ringing studies calls for all sorts of non-breedingareas and by hunting. Two other migratory survivalanalyses, even though the same wealth makes species,both of them curlews,are in even graverdanger the tasks ahead quite daunting. and are considered critical. The was hunted close to extinction in the last century, and was not As an example of another nice attempt at an integrationof seen for several decades until two birds were encountered existingbiological knowledge, we would like to mention in Texas in 1945. Eskimo Curlewswere reported in 24 of Ted Miller's reviews of the vocal communication behaviour the next 40 years, but the observednumbers were never of shorebirds(Miller 1992, 1995). The lists of what is largerthan 23 individuals. Still, the Eskimo Curlew may known, and what isn't, should now be extended to studies lingeron to this very day as suggestedby the sporadic of the kinds of messagestransmitted by shorebirds. If a sightings.The reasonsfor the demiseof the Slender- call is associatedwith a particularcommunicative function billed Curlews certainly includethe heavy hunting that should be quite general (e.g., a chick calling a parent pressureon its Mediterraneanwintering grounds, but this to brooding, or vice versa) and if the particular signal processmay havebeen aggravated by serioushabitat appears missingin the describedrepertoire of a shorebird loss both on the breedinggrounds and in the winter range. species,we better go out and look for it! Althoughthey are observedevery winter in very small numbers,the speciesis likelyto involvefewer than 100 individualsand is seriouslyendangered.

29 CONCLUSION del Hoyo,J., Elliott,A. & Sargital,J. (eds.) 1996. Handbookof the Birdsof the Wodd,Vol. 3. Hoatzinto Auks. LynxEdicions, Barcelona. There is enormous scope to do importantwork for the curious, travelling and writing naturalist and the hard Goss-Custard,J.D. (ed.) 1996. The Oystercatcher.From desk-boundworker! The tables presentedin the brief individualsto populations.Oxford University Press, Oxford. overview,and the more extensivetexts on ploversand Miller,E.H. 1992. Acousticsignals of shorebirds.A surveyand sandpipersin the Handbookof the Birdsof the World, Vol. reviewof publishedinformation. Royal British Columbia 3 (del Hoyo et al. 1996) should direct your interestfurther. MuseumTechnical Report, Victoria, B.C. Miller,E.H. 1995. Soundsof shorebirds:opportunities for amateurs andan updateof publishedinformation. Wader Study Group ACKNOWLEDGEMENTS Bull. 78:18-22.

Miskelly,C.M. 1990. Breedingsystems of NewZealand Snipe We thank Josep del Hoyo, Andy Elliottand Sheila Hardy Coenoryphaaucklandica and ChathamIsland Snipe C. pusilia, for their guidance,trust, patience and encouragement are they food limited? Ibis 132: 366-379. duringthe productionof our handbook-contributionsand their permissionto reproducethe informativefamily Myers,J.P. 1981. Cross-seasonalinteractions in the evolutionof "trees" in black-and-white. We thank Dick Visser for sandpipersocial systems. Behav. EcoL SociobioL8: 195-202. processingthe colour figures from the Handbook,and Piersma,T. & Wiersma,P. 1996. FamilyCharadriidae (plovers). In YvonneVerkuil and Petra de Goeij for much help and del Hoyo,J., Elliott,A. & Sargital,J. (eds.) Handbookof the niceties over the long period of Handbook-work. This is Birdsof the Wodd,Vol. 3. Hoatzinto Auks:384-442. Lynx NIOZ-publication 3029. Edicions, Barcelona. Piersma,T., van Gils, J. & Wiersma,P. 1996. FamilyScolopacidae (sandpipers,snipes and phalaropes).In: del Hoyo,J., Elliott, REFERENCES A. & Sargital,J. (eds.) Handbookof the Birdsof the World, Vol. 3. Hoatzinto Auks:'I,'I.'I. 533. LynxEdicions, Barcelona. Boyd,H. 1962. Mortalityand fertilityof the EuropeanCharadrii. Ibis Reynolds,J.D. & Sz•kely,T. 1997. The evolutionof parentalcare in 104: 68-87. shorebirds:life histories,ecology and sexualselection. Behav. Eco/.: in press. Collar,N.J., Crosby,M.J. & Stattersfield,A.J. 1994. Birds to watch 2: the wodd list of threatened birds. BirdLife Conservation Sz•kely,T. & Reynolds,J.D. 1995. Evolutionarytransitions in Series4. BirdLifeInternational, Cambridge. parentalcare in shorebirds. Proc. R. Soc. Lond. B 262:57-64

Appendix 1. Summarylist of the shorebirdspecies belonging to Charadriidaeand Scolopacidaethat were researchedfor the Handbookof the Birdsof the World,Vol. 3, includingnotes on their distribution and conservationstatus (according to designationsby BirdLifeInternational, Cambridge),the minimumnumber of publicationsreporting on aspectsof the speciesbiology, and the extentof knowledgeabout population size, demographicstructure, reproductive biology, migration systems and food and foraging. Publication record gives approximate number of topical publicationson the speciespublished before late 1995, where 1-10=A (not or poorlystudied), 11-20=B (a littleattention), 21-30=C (some more attention),31-40=D (well studied), 41-50=E (very well studied) and >50=F (extensivelystudied). Breeding regions: Eu=Europe (including Greenland),Af=Africa, As=Asia, Au=Australia, Oc=Oceania, NA=North America, SA=South America. Status: NGT=not globally threatened, N.Threat.=near-threatened,Vuln.=Vulnerable, Endang.=Endangered (see text). Near-threatenedto Endangeredspecies are printedin bold. The questionsare answeredwith a clear No, a clear Yes, or + = some information.

Englishname Latinname Public- Breeding Status Popu- Demo- Repro- Migra- Food& ation region lation graphic ductive tion fora- record size structure biology system ging known? known? known? known? known'• NorthernLapwing Vane#usvane#us F Eu,As NGT + Yes Yes Yes + Long-toedLapwing Vane#uscrassirostrls A Af NGT No No Yes No + Blacksmith Plover Vane#us armatus B Af NGT No No Yes + + Spur-wingedPlover Vane#usspinosus B Af, As NGT No No Yes + + RiverLapwing Vane#usduvaucelii A As NGT + No + No No Black-headed Vane#us tectus A Af NGT No No Yes + + Lapwing Yellow-wattled Vane//us malabarlcus A As NGT No No Yes No + Lapwing White-headed Vane#usa/biceps A Af NGT No No Yes No + Lapwing Lesser Black- Vane#uslugubrls B Af NGT No No Yes + No wingedLapwing Greater BI.-winged Vane#usmelanopterus B Af NGT No No Yes + No Lapwing CrownedLapwing •/anelluscoronatus B Af NGT No No Yes + + AfricanWattled Vane#ussenegal/us A Af NGT No No Yes + + Lapwing

30 Englishname Latin name Public- Breeding Status Popu- Demo- Repro- Migra- Food & ation region lation graphic ductive tion fora- record size structure biology system ging known? known? known? known? known? Spot-breasted Af NGT No No No No No Lapwing melanocephalus Brown-chested Vane#ussuperciliosus Af NGT No No No Lapwing Grey-headed Vane#us cinereus As NGT No No No Lapwing Red-wattled Vane#us indicus As NGT No No No Lapwing Javanese Wattled Vanellusmacropterus As Extinct Yes No No No Lapwing Banded Lapwing Vane#us tricolor Au NGT Yes No +_ MaskedLapwing Vane#us miles Au,Oc NGT Yes Yes Sociable Lapwing Vanellus gregarius Eu,As Vuln. No + White-tailed Vane#us leucurus Eu,As NGT No No _+ No Lapwing Pied Lapwing Vane#us cayanus SA NGT No No No No No SouthernLapwing Vanellus chilensis SA NGT No No No AndeanLapwing Vane#usresplendens SA NGT No No No No Red-kneed Dotterel Erythrogonyscinctus Au NGT Yes No No Eurasian Golden Pluvialisapricaria Eu,As NGT No Yes Yes Plover Pacific Golden Pluvialis fulva As,NA NGT No Yes Plover American Golden Pluvialis dominica NA NGT No Yes Plover Grey Plover Pluvialissquatarola NA,Eu,As NGT Yes ----. Yes Yes Yes Red -breasted Charadrius obscurus Oc Endang. Yes Yes Yes No Plover Common Ringed Charadrius hiaticula NA,Eu,As NGT ----. Yes Yes Yes Yes Plover Semipalmated Charadrius NA NGT No Plover semipalmatus Long-billedPlover Charadriusplacidus As NGT No No No No LittleRinged Plover Charadrius dubius Eu,As NGT No Yes No Wilson's Plover Charadrius wilsonia NA,SA NGT No No Yes Killdeer Charadrius vociferus NA,SA NGT No Yes No Piping Plover Charadrius melodus NA Vuln. Yes Yes Yes Yes Black-banded Charadrius Af Vuln. No No No No Plover thoracicus Kittlitz's Plover Charadriuspecuarius Af NGT No Yes St Helena Plover Charadrius Af Endang. Yes No sanctaehelenae Three-banded Charadrius tricollaris Af NGT No No No No Plover Forbes's Plover Charadrius forbesi A Af NGT No No No White-fronted Charadriusmarginatus C Af NGT No No Plover Kentish Plover Charadrius E xpt Au,Oc NGT Yes Yes Yes alexandrinus Javan Plover Charadriusjavanicus A As NGT No No No No Red-cappedPlover Charadriusruficapillus B Au NGT Yes No Yes No MalaysianPlover Charadriusperonii A As NGT No No ----. No No Chestnut-banded Charadriuspallidus B Af NGT No No No No Plover Collared Plover Charadrius collaris B NA,SA NGT No No No Puna Plover Charadrius alticola A SA NGT No No No No No Two-banded Plover Charad#us falklandicus B SA NGT No No No Double-banded Charadrius bicinctus C Oc N GT Yes Yes Yes Plover Lesser Sandplover Charadriusmongolus C As NGT No ----. Greater Sandplover Charadrius leschenaull• B As NGT No ----. Caspian Plover Charadrius asiaticus B As NGT No Charadrius veredus B As NGT Yes No No No Charadriusmorinellus E Eu,As NGT Yes Rufous-chested Charadfius modestus A SA NGT No No Dotterel Mountain Plover Charadrius montanus C NA Vuln. Yes Yes Yes Yes Hooded Plover Charadrius rubricollis C Au Vuln. Yes Yes Yes

31 Englishname Latin name Public- Breeding Status Popu- Demo- Repro- Migra- Food & ation region lation graphic ductive tion fora- record size structure biology system ging known? known? known? known? known'• Shore Plover Charadrius Oc Endang Yes Yes Yes Yes novaeseelandiae Black-fronted Elseyomismelanops Au,Oc NGT Yes Dotterel Inland Dotterel Peltohyasaustralis Au NGT Yes No + _+ Wrybill Anarhynchus Vuln. Yes Yes Yes Yes Yes frontalis Diademed Plover Phegomis mitchelli SA N.Threat No No No No Tawny-throated Oreopholusruficollis SA NGT No No No No Dotterel Magellanic Plover Pluvianellus socialis SA N.Threat + No + + Scolopaxrusticola Eu,As NGT No Yes Yes Yes Amami Woodcock $colopax mira As Vuln. No No No No No Rufous Woodcock Scolopaxsaturata As NGT No No No No No Sulawesi Scolopax celebensis As N.Threat No No No No No Woodcock Moluccan Scolopax rochussenii As Vuln. No No No No No Woodcock American Scolopaxminor NA NGT No Yes Yes Yes Yes Woodcock Chatham Snipe Coenocorypha pusilia Oc Vuln. Yes No Yes Yes Coenocorypha Oc N.Threat Yes No Yes Yes Snipe aucklandica Lymnocryptes minimus Eu,As NGT No No Gallinago solitaria As Vuln. No No No No No Latham's Snipe Gallinago hardwickill As N.Threat Yes No + + + Wood Snipe Gallinago nemoricola As Vuln. No No No No No PintailSnipe Gallinagostenura As NGT No No Yes Swinhoe'sSnipe Gallinagomegala As NGT No No No AfricanSnipe Gallinagonig#pennis Af NGT No No No No MadagascarSnipe Gallinagomacrodactyla Af NGT No No No No No Gallinago media Eu N.Threat + Yes CommonSnipe Gallinagogallinago NA,Eu,As NGT + Yes Yes Yes Yes South American Gallinagoparaguaiae SA NGT No No No No Snipe NobleSnipe Gallinagonob#is SA NGT No No No No No GiantSnipe Gallinagoundulata SA NGT No No No No No Gallinago stricklandii SA N.Threat No No No No No AndeanSnipe Gallinagojamesoni SA NGT No No No No No Imperial Snipe Gallinago imperialis SA N.Threat No No No No No Short-billed Limnodromusg#seus NA NGT No No Dowitcher Long-billed Umnodromus NA NGT No Dowitcher scolopaceus Limnodromus As N.Threat semipalmatus Black-tailed Godwit Lirnosa limosa Eu,As NGT Yes Yes Yes Yes Yes Limosa haemastica NA N.Threat + No Yes + + Bar-tailed Godwit Limosalapponica Eu,As NGT Yes No ----. Yes Yes Limosa fedoa NA NGT Yes No ----. ----. _+ Numenius minutus As NGT ----. No ----. ----. + Eskimo Curlew Numenius borealis NA Critical Yes No No + Whimbrel Numeniusphaeopus NA,Eu,As NGT Yes Yes Yes Bristle-thighed Numenius tahltiensis NA Vuln. Yes Yes Yes + Curlew Slender-billed Numenius As Critical Yes No No No Curlew tenuirostris Numenius arquata Eu,As NGT Yes Yes Yes Yes Far Eastern Numenius As N.Threat Yes No + _+ Curlew madagascariensis Long-billedCurlew Numenius ame#canus NA NGT No Yes Yes _+ _+ UplandSandpiper BartramiaIongicauda NA NGT No No Yes + + SpottedRedshank T#nga erythropus Eu,As NGT No _+ _+ Common Redshank Tringatotanus Eu,As NGT Yes Yes Yes Yes Tringastagnab?is Eu,As NGT No _+ _+ Common T#nga nebula#a Eu,As NGT Yes Yes + _+ Greenshank

32 Englishname Latin name Public- Breeding Status Popu- Demo- Repro- Migra- Food & ation region lation graphic ductive tion fora- record size structure biology system ging known? known? known? known? known? Nordmann's Tringa guttifer B As Endang. No Greenshank GreaterYellowlegs Tringamelanoleuca A NA NGT Yes No + Yes + LesserYellowlegs Tringafiavipes B NA NGT + No Yes Yes + Tringaochropus B Eu,As NGT No No Yes + + SolitarySandpiper Tringasolitaria A NA NGT No No No + + Tringaglareola B Eu,As NGT No No + + + Terek'sSandpiper Xenuscinereus B Eu,As NGT + No + + + CommonSandpiper Actirishypoleucos D Eu,As NGT No Yes Yes + + SpottedSandpiper Actitismacularia C NA NGT No Yes Yes + + Grey-tailedTattler Heteroscelusbrevipes A As NGT No No + + _+ Wandering Heteroscelusincanus A As,NA NGT + No + + + Catoptrophorus B NA NGT No No + + + semipalmatus White-winged Prosobonialeucoptera A Oc Extinct Yes No No No No Sandpiper MooreaSandpiper Prosobonia ellisi A Oc Extinct Yes No No No No Tuamotu Prosobonia B Oc Endang. Sandpiper cancellata RuddyTurnstone Arenariainterpres E NA,Eu,As NGT + Yes Yes Yes Black Arenaria A NA NGT Yes No _+ + melanocephala Aphriza virgata B NA NGT Yes No +_ + Calidris tenuirostds C As NGT Yes Yes Yes Red Knot Calidriscanutus F NA,E u,As NGT Yes Yes Yes Yes Caliddsalba E NA,Eu,As NGT _+ Yes Yes Yes Semipalmated Calidrispusilia D NA NGT Yes Yes Yes Yes + Sandpiper Calidrismauri D As,NA NGT + Yes Yes Yes + Red-necked Calidrisruficollis C As NGT Yes Yes Yes Yes + Calidrisminuta B Eu,As NGT + No Yes Yes + Temminck's Stint Calidristemminckii B Eu,As NGT No Yes Yes + No Long-toedStint Calidrissubminuta B As NGT No No Yes + No LeastSandpiper Calidrisminutilla C NA NGT No No Yes Yes + White-rumped Calidrisfuscicollis C NA NGT + No Yes Yes + Sandpiper Baird'sSandpiper Calidrisbairdii B As,NA NGT + No Yes Yes + PectoralSandpiper Calidrismelanotos B As,NA NGT No No Yes + + Sharp-tailed Calidrisacuminata B As NGT Yes No Yes + + Sandpiper CurlewSandpiper Calidrisferruginea E As NGT Yes + Yes Yes + PurpleSandpiper Calidrismaritima C NA,Eu NGT + + Yes Yes Yes RockSandpiper Calidrisptilocnemis B As,NA NGT No No Yes + + Dunlin Calidrisalpina F NA,Eu,As NGT Yes Yes Yes Yes Yes Spoon-billed Eurynorhynchus C As Vuln. Yes Yes Yes + + Sandpiper pygmaeus Broad-billed Limicolafalcinellus C Eu,As ' Vuln. + No Yes + + Sandpiper StiltSandpiper Micropalama B NA NGT + No Yes + + himantopus Buff-breasted Tryngitessubruficollis C As,NA NGT + No Yes + + Sandpiper Ruff Philomachuspugnax F Eu,As NGT + + Yes Yes + Wilson'sPhalarope $teganopustricolor E NA NGT Yes + Yes Yes Yes Red-necked PhalaropusIobatus E NA,Eu,As NGT + Yes Yes Yes + Phalaropusfulicaria D NA,EuIAs NGT No No Yes + +

33