Bulletin 39 - Yale Peabody Museum of Natural History - Yale University [PDF]

A Comparative Study of the Egg White Proteins of Non-Passerine Birds

CHARLES G. SIBLEY and JON E. AHLQUIST Peabody Museum of Natural History and Department of Biology, Yale University

BULLETIN 39 • 1 DECEMBER 1972 PEABODY MUSEUM OF NATURAL HISTORY YALE UNIVERSITY NEW HAVEN, CONNECTICUT

,

Bulletins published by the Peabody Museum of Natural History, Yale University, are numbered consecutively as independent monographs and appear at irregular intervals. Shorter papers are published at frequent intervals in the Peabody Museum Postilla series.

T h e Peabody Museum Bulletin incorporates the Bulletin of the Bingham Oceanographic Collection, which ceased independent publication after Vol. 19, Article 2 (1967).

Publications Committee:

John H. Ostrom, Chairman Theodore Delevoryas Willard D. H a r t m a n Keith S. Thomson Thomas Uzzell Charles G. Sibley, ex officio

Editor, Peabody Museum Publications: Zelda Edelson Special Editor: Jeanne E. Remington, editor this volume

Communications concerning purchase or exchange of publications should be addressed to the Publications Office, Peabody Museum of Natural History, Yale University, New Haven, Connecticut 06520, U.S.A.

Printed in the United States of America

CONTENTS LIST OF FIGURES

v

L I S T O F TABLES

vi

ABSTRACTS (ENGLISH, GERMAN, FRENCH, RUSSIAN)

1

INTRODUCTION A Chronological Survey of the Classification of Birds Summary Some of the Principal Characters Used in the Classification of Birds 1. Palatal structure 2. Pelvic musculature 3. T h e deep plantar (or deep flexor) tendons 4. Convolutions of the intestinal tract f 5. The carotid arteries 6. T h e fifth secondary Summary The Egg White Proteins Lysozyme Conalbumins Component 18 Globulins Ovomucoid Ovalbumin Prealbumins Mucins Materials and Methods

4 5 13 13 14 16 17 20 22 25 28 29 30 30 32 32 32 33 34 35 35

O R D E R S P H E N I S C I F O R M E S (Penguins) Introduction Historical Review of the Classification Summary T h e Egg White Protein Evidence Conclusions

36 36 36 42 42 43

T H E R A T I T E S (Ostriches, Rheas, Cassowaries, Emus), K I W I S AND T I N A M O U S Introduction Historical Review of the Classification Summary T h e Egg White Protein Evidence Co/iclusions

44 44 44 50 50 52

ORDERS GAVIIFORMES AND PODICIPEDIFORMES (Loons and Grebes) Introduction

53 53

Historical Review of the Classification Summary The Egg White Protein Evidence Conclusions

54 57 57 58

ORDER PROCELLARIIFORMES (Albatrosses, Shearwaters, and Petrels) . . Introduction Historical Review of the Classification Summary The Egg White Protein Evidence Conclusions

59 59 59 63 63 64

ORDER PELECANIFORMES (Tropicbirds, Pelicans, Frigatebirds, and Allies) Introduction Historical Review of the Classification Summary The Egg White Protein Evidence Conclusions

65 65 66 69 70 71

ORDER CICONIIFORMES (Herons, Storks, and Allies) Introduction Historical Review of the Classification A. Herons, Storks, Ibises Summary for Ciconiae B. Flamingos Summary for Phoenicopteri The Egg White Protein Evidence Conclusions

72 72 73 73 79 80 84 84 86

ORDER ANSERIFORMES (Screamers, Ducks, Geese, and Swans) Introduction Historical Review of the Classification Summary The Egg White Protein Evidence Conclusions

87 87 87 92 93 94

ORDER FALCONIFORMES (Vultures, Hawks, and Falcons) Introduction Historical Review of the Classification Summary The Egg White Protein Evidence Conclusions

95 95 96 102 103 105

ORDER GALLIFORMES (Megapodes, Curassows, Pheasants, and Allies) . . . Introduction Historical Review of the Classification Summary The Egg White Protein Evidence Conclusions

106 106 107 Ill Ill 113

ORDER GRUIFORMES (Cranes, Rails, and Allies) Introduction

114 114

Historical Review of the Classification Summary The Egg White Protein Evidence Conclusions ORDER CHARADRIIFORMES (Shorebirds, Gulls, and Auks) Introduction Historical Review of the Classification Summary The Egg White Protein Evidence Conclusions

115 ,. 124 124 126 . 127 127 128 140 141 143

ORDER COLUMBIFORMES (Sandgrouse, Pigeons, and Doves) Introduction Historical Review of the Classification Summary The Egg White Protein Evidence Conclusions

145 145 146 156 157 158

ORDER PSITTACIFORMES (Lories, Parrots, and Macaws) Introduction Historical Review of the Classification Summary The Egg White Protein Evidence Conclusions

159 159 160 166 166 167

ORDER CUCULIFORMES (Turacos, Cuckoos, and Hoatzin) A. Cuculiformes except Opisthocomus Introduction Historical Review of the Classification Summary The Egg White Protein Evidence . . • • Conclusions _, B. Opisthocomus hoazin (Hoatzin) Introduction Historical Review of the Classification Summary The Egg White Protein Evidence Conclusions

168 168 168 170 175 175 176 176 176 177 181 . 181 182

ORDER STRIGIFORMES (Owls) Introduction Historical Review of the Classification Summary The Egg White Protein Evidence Conclusions

183 183 184 189 190 190

ORDER CAPRIMULGIFORMES (Oilbirds, Goatsuckers, and Allies) Introduction Historical Review of the Classification Summary

191 191 192 195

The Egg White Protein Evidence Conclusions

196 197

ORDER APODIFORMES (Swifts and Hummingbirds) Introduction Historical Review of the Classification Summary The Egg White Protein Evidence Conclusions ,

198 198 198 205 205 206

ORDER COLIIFORMES (Colies) Introduction Historical Review of the Classification Summary The Egg White Protein Evidence Conclusions

207 207 208 211 212 212

ORDER TROGONIFORMES (Trogons) Introduction Historical Review of the Classification Summary The Egg White Protein Evidence Conclusions

213 213 213 217 217 218

ORDER CORACIIFORMES (Kingfishers, Bee-eaters, Rollers, and Hornbills) 219 Introduction 219 Historical Review of the Classification 220 Summary 228 The Egg White Protein Evidence 228 Conclusions 230 ORDER PICIFORMES (Jacamars, Barbets, Toucans, and Woodpeckers) . . . Introduction Historical Review of the Classification Summary The Egg White Protein Evidence Conclusions

231 231 232 238 238 239

PROBABILITIES AND POSSIBILITIES

240

ACKNOWLEDGMENTS

242

EPILOGUE

245

LITERATURE CITED

246

LIST OF FIGURES 1. Diagram of the starch gel electrophoretic pattern of the egg white proteins of a non-passerine bird 29 Starch gel electrophoretic patterns of: at back of book 2. The Spheniscidae, Struthionidae, Rheidae, Casuariidae, Dromaiidae, and Apterygidae (part) 3. The Apterygidae (part), Tinamidae, and Gaviidae 4. The Podicipedidae and Diomedeidae 5. The Procellariidae, Hydrobatidae, and Pelecanoididae (part) 6. The Pelecanoididae (part), Phaethontidae, Pelecanidae, Sulidae, and Phalacrocoracidae (part) 7. The Phalacrocoracidae (part), Anhingidae, Fregatidae, and Ardeidae (part) 8. The Ardeidae (part) 9. The Ardeidae (part), Scopidae, Ciconiidae, and Threskiornithidae (part) 10. The Threskiornithidae (part), Phoenicopteridae, Anhimidae, and Anatidae (part) 11. The Anatidae (part) 12. The Anatidae (part) and Cathartidae 13. The Pandionidae and Accipitridae (part) 14. The Accipitridae (part) 15. The Accipitridae (part), Sagittariidae, andFalconidae 16. The Megapodiidae, Cracidae, and Phasianidae (part) 17. The Phasianidae (part) 18. The Phasianidae (part), Turnicidae, Gruidae, Aramidae, Psophiidae, and Rallidae (part) 19. The Rallidae (part) and Rhynochetidae 20. The Eurypygidae, Cariamidae, Otididae, Jacanidae, Rostratulidae, Haematopodidae, and Charadriidae (part) 21. The Charadriidae (part) and Scolopacidae (part) 22. The Scolopacidae (part), Recurvirostridae, Phalaropodidae, Burhinidae, and Glareolidae 23. The Thinocoridae, Chionididae, Stercorariidae, and Laridae (part) 24. The Laridae (part) 25. The Laridae (part) and Alcidae 26. The Pteroclidae and Columbidae (part) 27. The Columbidae (part) 28. The Columbidae (part) 29. The Psittacidae (part) 30. The Psittacidae (part) 31. The Musophagidae and Cuculidae (part) 32. The Cuculidae (part), Tytonidae, and Strigidae (part) 33. The Strigidae (part), Steatornithidae, Podargidae, Nyctibiidae, Aegothelidae, and Caprimulgidae 34. The Apodidae, Hemiprocnidae, and Trochilidae (part) 35. The Trochilidae (part), Coliidae, Trogonidae, and Alcedinidae 36. The Todidae, Momotidae, Meropidae, Coraciidae, Upupidae, and Phoeniculidae 37. The Bucerotidae, Galbulidae, Bucconidae, Capitonidae, Indicatoridae, and Picidae

LIST OF TABLES 1. 2. 3. 4.

Principal anatomical differences between loons and grebes Principal anatomical characters of Gruiformes Differences between the Cuculidae and Musophagidae . . Characters of Tytonidae and Strigidae

53 116 169 184

ABSTRACT The starch gel electrophoretic patterns of the egg white proteins of 816 species of non-passerine birds were studied. Evidence of family-level systematic relationships was sought and compared with published data on anatomy, behavior, hybridization, parasites, the fossil record, biochemistry, biogeography and egg shell structure. In many cases the previously accepted classification was supported; in others, questions were raised by the protein data. The history of avian classification is outlined, and the principal characters that have been used in the classification of birds are evaluated with the aid of published data. It was concluded that: 1) the large ratites are monophyletic and evolved from a flying ancestor; 2) Pelecanoides is a procellariid; 3) Cochlearius is closest to Nycticorax; 4) the Phoenicopteridae are closer to the ciconiiforms than to the anseriforms, with a common ancestor for all three; 5) Opisthocomus is a cuculiform; 6) the Alcidae are closely related to the other charadriiforms; 7) Tyto is closely related to the Strigidae; 8) the Picidae and Capitonidae are closely allied. It is probable that: 1) the nearest relatives of the Sphenisciformes are the Procellariiformes; 2) the Tinamiformes are closest to the Galliformes; 3) Sula, Pelecanus, Phalacrocorax, and Anhinga are closely allied; 4) Scopus is a ciconiid; 5) the New World Cathartidae are closer to the other diurnal raptors than to the Ciconiidae; 6) the Gruidae, Aramidae, Rallidae, Eurypygidae, Heliornithidae, and Turnicidae form a natural assemblage; 7) among caprimulgiforms two natural groups exist, one of the Aegothelidae and Podargidae, the other of the Caprimulgidae, Nyctibiidae, and Steatornithidae; 8) the Apodidae and Trochilidae are more closely allied than either is to any other group; 9) the Coraciiformes of Wetmore are polyphyletic; 10) the Alcedinidae and Todidae are closely allied. Further conclusions on classification and relationships are given and ranked as possible or as improbable.

R£SUM£ Les profils electrophoretiques sur gel d'amidon des proteines de blanc d'oeuf de 816 especes de Non-Passereaux ont 6te etudies. Des evidences des relations systematiques au niveau des families ont etc* recherch£es et ces resultats compares aux donnees pubises concernant l'anatomie, le comportement, l'hybridation, les parasites, les fossiles, la biochimie, la biogeographie et la structure des coquilles d'oeufs. Dans de nombreux cas, la classification acceptee au pr^alable a £te soutenue; mais dans les autres cas les donnees proteiques ont pose des questions. L'histoire de la classification des oiseaux est esquissee et la validite des principaux caracteres utilises pour la classification avienne a et£ evaluee a Paide des donnees publiees. 1

2

PEABODY MUSEUM BULLETIN 39

Les suivantes conclusions sont considerees comme "tres probables": 1) les grands Ratites sont monophyletiques et ont evolues a partir d'un ancetre volant; 2) Pelecanoides est un procellaride; 3) Cochlearius est Foiseau le plus proche de Nycticorax; 4) les Phoenicopterides sont plus proches des Ciconiiformes que des Anseriformes, avec un ancetre commun aux trois; 5) Opisthocomus est un cuculiforme; 6) les Alcides sont de tres proches parents des autres Charadriiformes; 7) Tyto est etroitment apparente aux Strigides; les Picides et les Capitonides sont etroitment allies. Les relations suivantes sont "probables" : 1) les plus proches parents des Sphenisciformes sont les Procellariiformes; 2) les Galliformes sont les oiseaux les plus proches des Tinamiformes; 3) Sula, Pelecanus, Phalacrocorax, et Anhinga sont etroitment allies; 4) Scopus est un ciconiide; 5) les Cathartides du Nouveau Monde sont plus proches des autres rapaces diurnes que des Ciconiides; 6) les Gruides, Aramid£s, Eurypygides, Heliornithides et Turnicides forment un assemblage naturel; 7) parmi les Caprimulgiformes deux groupes naturels sont distingues, les Aegothelides et Podargides d'une part, les Caprimulgides, Nyctibiides et Steatornithides d'autre part; 8) les Apodides et les Trochilides sont plus proches entre eux qu'ils ne le sont separement de n'importe quel autre groupe d'oiseaux; 9) les Coraciiformes definis par Wetmore sont polyphyletiques; 10) les Alcedinides et les Todides sont etroitment allies. D'autre conclusions sur la classification et sur des parentes sont mentionees et designees ou bien "possibles" ou bien "improbables".

ZUSAMMENFASSUNG Die am Starke-Gel durchgeforschten elektrophoretischen Muster der EiweissProteine von 816 Arten der Non-Passeres wurden untersucht. Beweise systematische Verwandtschaften auf dem Rang der Familien wurden gesucht und wurden mit ahnlichen veroffentlichten Beweise aus Anatomie, Verhaltens, Bastardierung, Parasiten, Palaontologie, Biochimie, Biogeographie, und Struktur der Eierschalen verglichen. In vielen Fallen wurden die angenommene Einteilung bestatigt; in anderen Fallen wurden Fragen von Seiten der Proteinangaben aufgeworfen. Die Geschichte der Einteilung der Vogel wurde skizziert, und die vorher in der Einteilung der Vogel benutzte Hauptmerkmale wurden untersucht, mit Hilfe der veroffentlichten Data. Zum Entschluss gekommen sind wir dass: 1) die grosse Flachbrustvogel sind monophyletisch; 2) Pelecanoides gehort zu den Procellariidae; 3) Cochlearius steht Nycticorax am nachsten; 4) die Phoenicopteridae stehen die Ciconiiformes naher als die Anseriformes, und alle Drei teilen einen Urahn; 5) Opisthocomus gehort zu den Cuculiformes; 6) die Alcidae sind nahe mit den anderen Charadriiformes verwandt; 7) Tyto und die Strigidae sind nahe miteinander verwandt; 8) die Picidae und die Capitonidae sind eng verbunden.

NON-PASSERINE EGG WHITE PROTEINS

3

Aller Wahrscheinlichkeit nach: 1) die Procellariiformes sind die nachste Verwandten der Sphenisciformes; 2) die Tinamiformes stehen die Galliformes am nachsten; 3) Sula, Pelecanus, Phalacrocorax, und Anhinga sind eng verbunden; 4) Scopus ist ein der Ciconiidae; 5) die Neuweltgeier (Cathartidae) sind naher mit den anderen taglichen Greifvogel verwandt als mit der Ciconiidae; 6) die Gruidae, die Aramidae, die Rallidae, die Eurypygidae, die Heliornithidae und die Turnicidae bilden eine natiirliche Versammlung; 7) die Caprimulgiformes bilden zwei natiirliche Gruppen, die Eine der Aegothelidae und Podargidae, und die Andere der Caprimulgidae, Nyctibiidae und Steatornithidae; 8) die Apodidae und Trochilidae stehen einander naher als jede steht zu irgendeiner anderer Gruppe; 9) die Coraciiformes nach Wetmore sind polyphyletisch; 10) die Alcedinidae und die Todidae sind eng verbunden. Weitere Entscheidungen iiber die Einteilung und die Verwandtschaften der Vogel werden beigebracht und in eine Reihe als "moglich" oder "unwahrscheinlieh" gestellt.

P E 3 K) M E EHJIH H3y^ieHH ie H3y*ieHHJi npoTenHOB B036y/i;HJiH Bonpocbi: ^aeTca KJiaccn$HKan;Hii nepHaTbix, a rjiaBHbie xapaKTepHbie ocoSeHHOCTH, Hcnojib30BaHHi)ie p a KjraccH$HKaii;HH nTHn;, oqeHHBaiOTCji npn noMonjH onySjiHKOBaHHHx ^aHHax. HTaK, SHJIH BHBe,n;eHBi 3aKJiK)qeHHa, ^TO 1) 6ojn>mHe HejieTaromne ETHI^H aBjiHioTca M0H0(|)HJieTH^ecKHMH H pa3BHJiHCb OT JieTaioniHx npe^KOB; 2) Pelecanoides 9T0 procellariid; 3) Cochlearius 6jiH3Ke Bcero K Nycticorax; 4) Phoenicopteridae SjiHJKe K $opMaM ciconii, *ieM K $opMaM anseri, c O6IH;HM npeflKOM /pa Bcex Tpex $opM; 5) Opisthocomus BTO $opMa cuculi; 6) Alcidae TecHO potfCTBemibi c flpyrHMH $opMaMH charadrii; 7) Tyto 6.3H3K0 pOflCTBeHHa c Strigidae; 8) Picidae H Capiton i d a e flBJUIIOTCJI TeCHO pOflCTBeHHblMH.

Bepoaraee Bcero, *ITO 1) 6jiH3KaftniHMH poflCTBeHHbiMH $opMaMH jpa Sphenisci aBJiJUOTca $opMH Procellarii; 2) $opMbi Tinami 6jiH3Ke Bcero K (J)opMaM Galli; 3) Sula, Pelecanus, Phalacrocorax H Anhinga 6JIH3KO poflCTBeHHbi; 4) Scopus BTO ciconiid; 5) Cathartidae HoBoro CBGTafijiHJKeKftpyrHM^HeBHbiM XHiipHKaM, qeM K Ciconiidae; 6) Gruidae, Aramidae, Rallidae, Eurypygidae, Heliornithidae H Turnicidae o6pa3yiOT HaTypajibHyio rpynny; 7) cpe^n (JwpM caprimulgi HMeioTCtf flBe ecTecTBeHHHe rpynnbi, OfljBa cocToain;aji H3 Aegothelidae H Podargidae, ftpyraH — H3 Caprimulgidae, Nyctibiidae H Steatornithidae; 8) Apodidae H Trochilidae ropa3,a;o Sraace poflCTBeHHH .zrpyr K flpyry, ^eM KajK^aa H3 HHX B OT^eJibHocTH K JIK)6OH flpyroft rpynne; 9) $opMbi Coracii no YfiTMopy (Wetmore) aBJijnoTca nojiH:-

Anas

platyrhynchos

A n a s pfatyrhy mhos

• # & • • • • •

A n a s acuta

FIG. 11. Starch gel electrophoretic patterns of the Anatidae ( p a r t ) .

#

Anas

georgica

Anas

punctata

Afihya

fuligula

Aythya

australis

Aythya

marila

&m& * # & * . • •

' & • ' • • •

Malacorhynchus

membranaceus

Somateria

mollissimo

Somaieria

speciabilis

Melaniita

nigra

Bucephala

Mergus

albeola

serrator

3 jamaicensis

i maccoa

Biziura

lobaia

Cathartes GOTO

€©ro§yps

atraius

FIG. 12. Starch gel electrophoretic patterns of the Anatidae (part) and Gathartidae,

Pcrncfion

haliaeius

Pernis apivorus Machaerhamphus

a/cinus

m ffotufs feucurus

f /anus

camulem

Icfinia

misisippiensis

Miiwm

migrans

Haliastur

m

sphenurus

Gyps africanus

Gyps coprofheres

Torgos fracheliofws

Circaefws galliens

$pif©f*nis cheela

Circws pyg org us

m

Circus cfefuginosus

Circus cyanews

FIG. 13. Starch gel electrophoretic patterns of the Pandionidae and Accipitridae (part).

'"5^

I Melierax

meioboies

W0' I Mccipiier melanoleucus

Kaupifalco

monogrammicus

Buteogallus '"!**T

anthracinus

Heterospizias

Parabuteo

mendionahs

vnkinctus

Buteo rufofuseus $

*

'^fBJIP

Buteo polyosoma

JSBE

Bufeo lineatus

m ipfe© swainsoni « Buteo

jamakensis

# Buteo bufeo 4;

4

Aquilo rapax Aquila chrysaefos Aquilo owdoK 4rf

Hieraeetus

pennatus

FIG, 14. Starch gel electrophoretic patterns of the Accipitridae ( p a r t ) ,

Lopfiaetus

Spizaetus

*p

occipitalis

cirrhafws

iPolemaetus belficosps

Sagittarius

serpentarius

!

Milwago chimangm

i

m

Po/i h/erax semiforqiiafiis

Folc©

naumanni

Falco

sparverius

#

Fafc© fffinuncu/us

Fctlc© cencfiroides

Folc© cofwwifaariws

Falco subbwfe©

Falco /ongipennis

Falco efeanorae

Falco biaritiicps

m

Falco

pemgrinus

FIG, 15. Starch gel electrophoretic patterns of the Accipitridae ( p a r t ) , Sagittariidae., and Falconidae,

Megapadius

•*

Alectura

Orialis

ij§|

i

lathami

cankollk

Penelope

Crax

superciiiaris

blumenbachii

Dendragapus

3 2

freycmet

Lagopus

obscurus

lagopus

Lagopus mutus

Tetrao urogallus

3^j&>'•*.'

Bonasa

umbellus

Centrocercus urophasianus

Ttagopan

w *

*

f

t

#

c W'

i

i f

temmincki

Gall us gall us

Lophura

swinhoei

Syrmaticus reeves!

Phasianus co/c/iicus

FIG. 18. Starch gel electrophoretic patterns of the Megapodiidae, Cracidae, and Phasianidae ( p a r t ) .

m

Chrysolophus pictus Chrysolophus

m

Polyplectron

*

.. *****

m

Powo

amherstiae raalacense

cristaius

Alecforis

mfa

Alectoris groeca

Alecforis borfacffo

Francolinus ercfcefii

i ^

% Perdix

perdix

iiKUBMBBMlli

Coturnix

coturnix

Exca/facforia cftinensls

Tropkoperdix #

•*m«.

*

charltonii

Rof/u/us rou/rou/

Callipepla

squomata

Lophortyx

douglasii

Lophortyx

californicus

*

FIG. 17. Starch gel electrophoretic patterns of the Phasianidae ( p a r t ) ,

*

•

m

Colinus

wirgimianus

Numida

meleagris

Meleagris

Turnix

gallopavo

sylvatica

Turnix

melanogaster

Turnix world

Bahama

pawonina

Aramus

guarauna

Pmphio

crepitans

Rallus longirostris

Rallus

limkola

Rallus caerulescens

Rallus

philippensis

Aramides

Crecopsis

cajanea

egregia

Limnocorax

flavirostra

*

FIG. 18, Starch gel electrophoretic patterns of the Phasianidae (part), Turnicidae, Gruidae, Aramidae, Psophiidae, and Rallidae (part).

Mm*

w

Porzcna

pusilla

Porzono Carolina

Laterallus

•

jamaicensis

Sarothrura

rufa

Poliolimnas

cinereus

Porphyriops

melanops

Amaurornis

phoenicurus

Gallinula

chloropus

Porphyrula

Porphyria

martinica

porphyria

Porphyria

madagascariensis

Fulica atra

Fulica cristata

Fulica

armillata

Fulica cornuta Rhynochetos

jubatus

FIG, 19. Starch gel electrophoretic patterns of the Rallidae (part) and Rhynochetidae,

Eurypyga

Cariama

helms

cristata

Otis tarda

Choriotis Icon

Lissotis melanogaster

Actophilornis

africana

Jacana spinosa

Rostratula

henghalensis

Nycficryphes

semicollam

Haematopus

ostralegus

Vaneilus

vaneilus

Vanellus or mat us

*

'•

Sll

^

^^-

.211

Voftellws coronafus

Vanellus chilensis

-*#$^|

gtt|

iflHBI

Pluviaiis

daminka

Pluvialis

squatarolo

FIG. 20, Starch gel electrophoretic patterns of the Eurypygidae, Cariamidae, Otididae, Jacanidae 3 Rostratulidae 5 Haematopodidae, and Gharadriidae ( p a r t ) .

Charadrius

hiatkula

Charadrius

dubius

Chamdrms

melonops

Eudromias

morinellus

Limosa limosa

Limosa

haemastica

Numenius

S

Tringa

phaeopus

tetanus

Catoptrophorus

Mrenaria

!

•

semipalmatus

interpres

GallinagG

gallinago

Gallinago

gallinagm

Gallinago

pormguaiae

«

Calidris canutus

t

Calidris

pusilla

'•Caiidris

maritima

FIG. 21. Starch gel electrophoretic patterns of the Gharadriidae (part) and Scolopacidae ( p a r t ) ,

4 t

i €1

Calidris

alpina

Limicola

falcinellus

Philomachus

pugnax

Himantopus

himantopus

Recurvirostfo

a¥oseffa

l©€t#F¥if©sfro

nowaehollandiae

Phalaropus

fulicarius

Phalaropus

lobatus

Burhinus capensis

Burhinus

oedicnemus

^T Burhinus bistriatus

Rhinoptilus

africanus

Peltohyas australis

Stiltia

Glareola

Glareola

Isabella

pratincole

cinema

FIG, 22. Starch gel electrophoretic patterns of the Scolopacidae (part), Recurvirostridae, Phaiaropodidae, Burhinidae, and Glareolidae.

m

Thinocorus erbigftyiofiffs

Affagls gcryi

Chionis

alba

« Stercorarius skua

Stercorarius

longicaudus

Stercorarius

parasiticus

tarus fridacfy/us

• #

*

Lams sabini

41

S

8 i

m

*

•

m

Larus

airkiUa

Larus pi pi xc an

Laruscanus

Larm

ridibundus

Lams marinus Larm

hyperboreus

afljk, # '

Larus thayeri

* Larus

argentatus

FIG. 23. Starch gel electrophoretic patterns of the Thinocoridae, .Ohionididae, Stercorariidae and Laridae (part).

Lams fuscus Lams

dominicanus

Lams crass/rosfr/s

tarws

delmwamnsis

Lams cirrocephalus Rynch&ps

T

#

nigra

Mnous stolidus

Aeons tenuirostris

Anous albus

AWOPS ceruleus

%

• v vr ?

Sterna

hybrido

Sterna

nilotica

Sterna

bergii

Sterna

fmrsteri

*&.. Sterna

dmugallii

Sterna

hirundo

FIG. 24. Starch gel electrophoretic patterns of the Laridae ( p a r t ) .

'^

^HHl •r;* ''4HMf|

* JP

;

Sterna nigra Sterna viffafa Ster no paradisaea Sterna lunata

«T

Sfer no fuscata

Sterna albifmns » Sterna lorota Plautus mile -

Lunda cirrhGta Fmterculm arctica *

Akm tmdm I

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