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BULLETIN OF
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THE BRITISH MUSEUM^ i 6 ju NI976
(NATURAL HISTORY)
ZOOLOGY
Vol. 27
1974
BRITISH MUSEUM (NATURAL HISTORY)
LONDON: 1976
DATES OF PUBLICATION OF THE PARTS
No. i . . . . . .13 June 1974
No. 2 -"i . . . . .18 July 1974
No. 3 19 July 1974
No. 4 . . . . .20 September 1974
No. 5 . . . . .23 December 1974
No. 6 . . . . .23 December 1974
No. 7 . . . . .24 December 1974
ISSN 0007-1498
Printed in Great Britain by John Wright and Sons Ltd. at The Stonebridge Press, Bristol BS4 5NU
CONTENTS
ZOOLOGY VOLUME 27
PAGE
No. I. Anatomy of head and neck in the Huia (Heteralocha acutirostris)
with comparative notes on other Callaeidae. By P. J. K. BURTON
(i PI.) . . i
No. 2. Miscellanea .......... 49
A new skink (Reptilia : Scincidae : genus Emoia) from the New
Hebrides. By LORD MEDWAY ...... 53
Scutocyamus parvus, a new genus and species of whale-louse
(Amphipoda : Cyamidae) ectoparasitic on the North Atlantic
white-beaked dolphin. By R. J. LINCOLN and D. E. HURLEY
(i PL) . . 59
Catalogue of the whale-lice (Crustacea : Amphipoda : Cyamidae)
in the collections of the British Museum (Natural History). By
R. J. LINCOLN and D. E. HURLEY 65
A review of Laephotis Thomas, 1901 (Chiroptera : Vespertilioni-
dae). By J. E. HILL 73
The genus Finmarchinella Swain, 1963 (Crustacea : Ostracoda)
and its species. By J. W. NEALE (2 Pis) .... 83
Euplotes rariseta sp. n. (Protozoa : Ciliatea). A new small
marine hypotrich. By C. R. CURDS, B. J. WEST and J. E.
DORAHY (i PL) 95
Observations on clonal cultures of Eugfypha acanthophora and
Euglypha strigosa (Testacea : Protozoa). By R. H. HEDLEY,
C. G. OGDEN and J. I. KRAFFT (2 Pis) 103
Descriptions of three species of Euplotes (Protozoa : Ciliatea).
By C. R. CURDS (i PL) 113
New records of bats from Southeastern Asia, with taxonomic
notes. By J. E. HILL 127
No. 3. The Haplochromis species (Pisces : Cichlidae) of Lake Rudolf, East
Africa. By P. H. GREENWOOD 139
No. 4. A review of Scotoecus Thomas, 1901 (Chiroptera : Vespertilionidae) .
By J. E. HILL 167
No. 5. Catalogue of the types of terrestrial isopods (Oniscoidea) in the
collections of the British Museum (Natural History) I. Superfamily
Pseudotracheata. By R. J. LINCOLN and J. P. ELLIS . . 189
No. 6. Campylaspis species (Crustacea : Cumacea) from the deep Atlantic.
By N. S. JONES -247
No. 7. A new family, genus and species of bat (Mammalia : Chiroptera)
from Thailand. By J. E. HILL 301
Index to Volume 27 . . . . . . 337
INDEX TO VOLUME 27
The page numbers of the principal references and the new taxonomic names are printed in bold type.
abbreviatus, Cyamus
abnormis, Gelsana .
abyssinicus, Microcercus .
Acaeroplastes .
Acanthoniscus
acanthophora, Euglypha .
Actoecia
Actoeciidae
aculeata, Campy laspis
acuminatus, Rhinolophus .
. 65
. 214
. 215
193
. 219
. 103-112, PI. I
. 218
. 191, 218
250, 253, 268-270
. 131
acuminatus, Rhinolophus acuminatus
131-132, 137
acuminatus acuminatus, Rhinolophus
131-132, 137
acuminatus audax, Rhinolophus . . 132
acuminatus calypso, Rhinolophus . -132
acuminatus circe, Rhinolophus . . .132
acuminatus sumatranus, Rhinolophus . 132
acustiserra, Porcellio .... 197
acutirostris, Heteralocha . . 1-48, i PI.
Adinda ....... 244
aediculatus, Euplotes . . . .124
aegaeum, Armadillidium .... 208
aegaeus, Porcellio . . . . . 197
aenigma, Diploexochus .... 223
aerarius, Diploexochus .... 224
affinis, Armadillo . . . . .219
affinis, Campylaspis. . 250, 254, 263, 277
affinis, Cubaris . . . . .219
affinis, Euplotes 113, 114, 116, 117-1 19, 120, 123
affinis tricirratus, Euplotes . 113, 119, 123
Agabiforius . . . . . -237
Agabiformius ...... 193
Agnara ....... 205
Akermania . . . . . .219
alassiense, Armadillidium . . 208, 212
alatus, Euplotes .... 97, 101
Alauda . . . . . . -41
alba, Campylaspis . . 250, 251, 256, 258
albanicum, Armadillidium . . . 208
alberti, Diploexochus . . . .224
albertianus, Haplochromis . . 150, 159
albescens, Diploexochus .... 224
albicornis, Lucasius .... 191, 197
albicornis, Mesarmadillo . . . -215
albicornis, Porcellio . . . . 197
albifrons, Armadillidium .... 208
albigula, Scotoecus . . 1 80, 1 8 1, 182, 183
albigula, Scotoecus hindei 180, 181, 182-i83
albinus, Porcellio . . . . .198
albipennis, Tura ..... 204
albirostris, Lagenorhynchus . . 59, 71
albofuscus, Scotoecus 169, 171, 172, 174-175,
176, 177, 186
albofuscus, Scotoecus albofuscus . 1 75-176
albofuscus, Scotophilus . . . 169, 175
albofuscus albofuscus, Scotoecus . 175-176
albofuscus cinnamomeus, Scotoecus . . 175
albofuscus woodi, Scotoecus . 175, 176-177
albomaculatus, Buddelundia . . .222
albomarginatus, Armadillo . . .219
albomarginatus, Porcellio . . . 198
albospinosus, Armadillo . . . .219
album, Armadillidium .... 208
alloniscus ...... 205
alluaudi, Porcellio . . . . .198
alticola, Diploexochus . . . .224
altimontis, Saidjahus .... 229
alveolata, Euglypha .... 104
ambitiosus, Armadillo . . . .229
ambitiosus, Spherillo .... 229
amblyoda, Campylaspis . . . 250, 251
Amblyrhamphus . . . . -37
americanum, Ethelum . . . -213
americanus, Mesarmadillo . . .213
amieti, Euplotes . . . . .124
amoenus, Porcellio ..... 198
Amphipoda . . . 59-64, I PI.; 65-72
ampullatus, Hyperoodon . . . -71
Anchicubaris . . . . . .219
anconanum, Armadillidium . . . 208
anconanus, Metoponorthus pruinosus . 197
aneityumensis, Emoia . . . 53-55, 56
Angara ....... 193
angolensis, Laephotis 73, 76-77, 7 8 - 79, 80, 81
angolensis, Laephotis wintoni . . -73
angularis, Campylaspis . . . 250, 251
angulata, Barentsovia . . 86, 87, 88-89,
91, 92, Pis. 1-2
angulata, Cythere .... 87, 88
angulata, Cythereis . . .83, 88
angulata, Finmarchinella . . 86, 87, 88-89,
90, 91, 92, Pis. 1-2
angulata, Hemicythere . . 86, 87, 88, 89
angulata, "Hemicytherinae" . . .88
angulata, Nereina . . 86, 87, 88, 90, 91
angusta, Tura . .204
angustulus, Porcellio .... 198
anna, Ciridops . 25, 28
annectans, Myotis . . 33
annulicornis, Cylisticus .... 192
antarctica, Campylaspis . . 250, 256, 299
Anthochaera . 34
apenninorum, Armadillidium . . . 208
338
INDEX
apenninorum, Trachelipus . . . 207
apenninorum, Tracheoniscus . . . 207
aperta, Campylaspis . . . 250, 253
apfelbecki, Armadillidium . . . 208
approximates, Metoponorthus . . .196
apulicus, Trachelipus .... 207
apulicus, Tracheoniscus .... 207
Arachnothera . . . . . .42
arcuata, Campylaspis . 250, 253, 265-266
arcuatus, Porcellio ..... 202
argentarium, Armadillidium . . . 208
argentarius, Acaeroplastes . . . 193
argentarius, Metoponorthus . . 193
Armadillidae . . 197, 219-232, 244-245
Armadillidiidae . 191, 208-213, 240-242
Armadillidium 208-212, 213, 221, 240, 241, 242
Armadillo . 208, 219-221, 222, 224, 225,
226, 227, 228, 229, 230, 231, 232, 244, 245
armatus, Benechinus . . . .213
artinii, Scotoecus . . . 180, 181, 182
artinii, Scotoecus hindei . . . .182
asifensis, Metoponorthus .... 197
asifensis, Metoponorthus sexfasciatus . 197
aspera, Campylaspis . . . 250, 255
aspheronicus, Euplotes . . . .124
assimile, Armadillidium .... 209
astriger, Armadillidium .... 208
astriger, Armadillo . . . .208
ater, Porcellio . . . . .198
aternanus, Trachelipus larii . . . 207
aternanus, Tracheoniscus larii . . . 207
aterrimus, Phoeniculus . . . 37, 45
Atlantidiidae . . . . .191
atrox, Phoniscus . . . 135-136, 137
audax, Rhinolophus aciminatus . . 132
aulacoeis, Campylaspis . . . 250, 252
Aulonocara . . . . . .160
auritus, Porcellio ..... 198
ausseli, Armadillo . . . . .219
australis, Eubalaena glaciolis . 67, 68, 69, 71
avium, Haplachromis . . . .159
badium, Armadillidium .... 208
badium siculorum, Armadillidium . . 208
Balaena . . . . . 67, 71
Balaenoptera . . . . 65, 66, 71
balaenopterae, Cyamus . . . 65-66, 71
Balantiopteryx . . . . -3*7
Balionycteris . . . . .128
Balloniscidae .... 191, 205
Balloniscus ...... 205
balstoni, Scoteinus . . . . . 173
balteatus, Euplotes ..... 101
balticus, Euplotes . . . . . 101
balticus, Porcellio ..... 207
balticus, Trachelipus .... 207
balticus burzenlandicus, Porcellio . . 207
balticus burzenlandicus, Trachelipus . . 207
barbertoni, Bethalus . . . .221
Barentsovia . .83, 84, 86, 87-92, Pis. 1-2
barentzovoensis, Barentsovia . 83, 86-88, 89,
90, 91, Pis. 1-2
barentzovoensis, Finmarchinella 83, 86-88, 89,
90, 91, Pis. 1-2
barentzovoensis, Nereina . . . 83, 86
barnardi, Bethalus ..... 222
barnardi, Cubaris . . . . .222
barroisi, Porcellio ..... 198
batesoni, Porcellio . . . . .198
benaci, Metoponorthus . . . .196
Benechinus . . . . . .213
bergomatius, Cylisticus plumbeus . .192
besi, Periscyphis . . . . .216
Bethalus .... 221-222, 245
bicarinata, Campylaspis . 250, 254, 287-28g
bicarinata, Merulana .... 228
bicarinata, Spherillo .... 228
bicolor, Exzaes . . . . .212
bicoloratus, Armadillo .... 229
bicoloratus, Periscyphis . . . .216
bicoloratus, Spherillo .... 229
bicornis, Mahehia . . . . . 195
biellensis, Cylisticus. .... 192
bifrons, Armadillo ..... 229
bifrons, Spherillo . . . . .229
bimaculatus, Hemichromis . . .141
binotatus, Buddelundia . . . .223
bipartitus, Buddelundia .... 223
bistriatus, Porcellio . . . .198
bisulcatus, Euplotes . . . 99, 101
bituberculatus, Armadillo . . .219
bituberculatus, Diploexochus . . .224
bizonatus, Periscyphops . . . .217
blattarius, Porcellio .... 198
bloyeti, Haplochromis . . 145, 162, 163
blythi, Rhinolophus . . . 131, 137
blythi, Rhinolophus blythi . . -131
blythi blythi, Rhinolophus . . -131
blythi calidus, Rhinolophus . . .131
blythi szechwanus, Rhinolophus . -131
bocki, Nesodillo ..... 228
bodenheimeri, Hemilepistus . . -194
bonetti, Campylaspis . . 250, 251, 261
boopis, Cyamus .... 66-67, 71
borelli, Plataoniscus .... 205
borellii, Alloniscus ..... 205
botswanae, Laephotis . 73, 76, 77-79, 80, 81
brachycephalus, Spherillo. . . .229
brachypterus, Philetor . . . . 134
brachypterus, Philetor brachypterus . . 135
brachypterus, Vespertilio . . -134
brachypterus brachypterus, Philetor . .135
brachypterus rohui, Philetor . . . 135
brachypterus verecundus, Philetor 134-135, 137
bredanensis, Steno .... 70, 71
brentanus, Trachelipus .... 207
brentanus, Tracheoniscus .... 207
brevicornis, Armadillo . . . .219
brevicornis, Campylaspis 250, 254, 270-271
brevipennis, Porcellio . . . .198
INDEX
339
brevis, Ignamba
brevis, Phalaba
buddelundi, Porcellio
Buddelundia .
bulbosa, Campylaspis
. 215
. 206
. 198
222-223
250, 254, 273-274, 289
bulgaricus, Trachelipus .... 207
bulgaricus, Tracheoniscus . . . . 207
bullatus, Haplochromis . . 159, 160, 161
Burton, P. J. K. . . . l- 4 8, i PI.
burzenlandicus, Porcellio balticus . . 207
burzenlandicus, Trochelipus balticus . . 207
Cacicus ..... 36, 37
cadornae, Pipistrellus .... 303
caelatum, Armadillidium .... 208
Caeroplastes . . . . . . 193
calidus, Rhinolophus blythi . . -131
caligans, Spherillo . . . . .229
Callaeas . . 3, 4, 6, 7, n, 12, 14, 22,
23. 26, 35-36, 44
Callaeidae .... 1-48, i PI.
callosus, Buddelundia .... 223
calmani, Porcellio ..... 198
calypso, Rhinolophus acuminatus . .132
Campy laspides . . . . .270
Campylaspis ..... 247-3 oo
Campylorhamphus . . . . -42
canaliculata, Campylaspis . 250, 252, 263
canaliculata, Merulana . . . .228
canaliculatus, Spherillo .... 228
canariensis, Armadillo .... 224
canariensis, Diploexochus. . . .224
canariensis, Porcellio . . . .198
caperata, Campylaspis . . . 250, 254,
279-280, 282
carinatus, Armadillo .... 222
carinatus, Bethalus ..... 222
carinatus, Hemiporcellio . . . -195
carniolense, Armadillidium . . . 208
Carolliinae . . . . . .319
carpathicus, Protracheoniscus saxonicus . 207
carthaginensis, Porcellio . . . .198
carunculata, Anthochaera . . -34
carunculatus, Creadion . 3, 4, 5, 6, 10, n, 34
cassida, Armadillo ..... 222
cassida, Bethalus . . . . .222
castor, Diploexochus . . . .224
catodon, Physeter . . . 66, 67, 70, 71
catodontis, Cyamus . . . 67, 71
caucasius, Cylisticus . . . .192
caudatus, Ourachaerus .... 232
cavernicola, Periscyphis . . .216
cayennensis, Porcellio . . . . 199
cela, Cacicus . . . . . -37
celsicauda, Diploexochus .... 224
ceti, Cyamus ..... 67, 71
charon, Euplotes . . 97, 113, 115, 123
chindeensis, Periscyphis . . .216
Chironax .... 127-129, 137
Chiroptera 73-82; 128-138; 167-188; 301-336
Cichlidae ..... 139-165
ciliata, Euglypha . . . . .104
Ciliatea . . 95-102, i PI.; 113-126, i PI.
cilicius, Metoponorthus . . . .196
cinctus, Armadillo ..... 220
cinctutus, Pyrgoniscus . . . .232
cinerascens, Buddelundia . . . 223
cinerea, Callaeas . . . 3, 4, 6, n, 12,
22, 23, 26, 35-36
cinerea, Creatophora . . . -37
cingendum, Armadillidium . . . 208
cingendum, Armadillidium maculatum . 208
cingulatus, Diploexochus .... 224
cinnamomomeus, Scotoecus albofuscus . 175
circe, Rhinolophus acuminatus . . . 132
Circoniscus . . . . . .218
circumdatus, Pipistrellus . . 132-134, *37
Ciridops ..... 25, 28
civilis, Periscyphis . . . . .216
clathrata nuda, Cythere . . . 86, 87, 88
clausus, Synarmadillo . . . .231
clavata, Campylaspis . . 250, 255, 292
Coelops ...... 320
cognata, Campylaspis 250, 254, 275, 277-278
cognatus, Porcellio . . . .199
Coleura ....... 323
collicolus, Porcellidium .... 206
collicolus, Porcellio ..... 206
collicolus, Porcellium .... 206
collinus, Armadillo . . . .220
coloratus, Diploexochus .... 224
compressa, Euglypha . . . .104
concinella, Normanicythere . . -85
concinnus, Scleropactes . . . .218
congener, Armadillo . . . .224
congener, Diploexochus . . . .224
conglobator, Armadillo . . . ^ . 220
conisaleus, Diploexochus . . . 224
contra, Sturnus . . . . -37
Contractus, Porcellio . . . . 199
Corcorax . . . . . .29
corcyraeum, Armadillidium . . . 209
cornutus, Rhinolophus . . . -131
cornutus pumilus, Rhinolophus . . -131
Corvus ....... 38
costata, Campylaspis . . 250, 253, 266
costata speciosa, Campylaspis . . . 266
coxalis, Metoponorthus .... 196
Graseonycteridae .... 301-336
Craseonycteris .... 301-336
crassus, Armadillo . . . . .224
crassus, Diploexochus .... 224
Creadion . . . 3, 4, 5, 6, 7, 10, n,
14. 34- 35. 36. 38. 44
Creatophora . . . . . -37
crenosus, Euplotes ..... 101
creper, Saidjahus ..... 229
crispa, Campylaspis. . . . 250, 253
cristatus, Euplotes . . . 99, 101, 121
340
INDEX
cristatus, Hemilepistus .... 194
cruentatus, Porcellio . . . . 199
Crustacea . 59-64, i PL; 65-72; 83-94,
2 Pis.; 189-246; 247-300
Cubaris . 219, 220, 221, 222, 225, 226, 231
Cumacea ..... 247-300
Curds, C. R. . 95-102, i PL; 113-126, i PL
curvicosta, Barentsovia
curvicosta, Finmarchinella
Cushmanidea
Cyamidae
Cyamus ....
cyanomelas, Rhinopomastus
Cylisticidae
Cylisticus
Cythere ....
Cythereis
. 83, 86, 87, 88,
89, 90-91, Pis. 1-2
. 83, 86, 87, 88,
89, 90-91, Pis. 1-2
. 86
59-64, i PL ; 65-72
59, 65-70, 71
. 42
191, 192
. 192
86, 87, 88
. 83, 84, 88
dalmatinus frascatensis, Orthometopon . 206
danae, Armadillo ..... 230
danae, Spherillo . . . . .220
davidi, Armadillidium .... 209
debueni, Porcellio ..... 199
decoratus, Spherillo . . . .229
decumanus, Psarocolius . . . -37
delphini, Isocyamus . . . 70, 71
dentifrons, Cylisticus . . . .192
depressus, Armadillo .... 222
depressus, Bethalus .... 222
Desertoniscus . . . . . .193
desertorum, Protracheoniscus . . . 206
Desmodontidae
diadaleos, Euplotes .
Diclidurinae .
Diclidurus
dimorphus, Porcellionides
Diploexochus .
disjunctus, Diploexochus .
dispar, Porcellio
dispersus, Spherillo . .
dollfusi, Eubelum
Dolphin, white-beaked
Dorahy, J. E.
dugesi, Armadillo
dugesi, Diploexochus
dumorum, Armadillo
dumorum, Venezillo
ecaudatus, Diploexochus
echinata, Campylaspis
egens, Armadillo
elbanum, Armadillidium
Ellis, J. P.
elongatus, Hemilepistus
Eluma .
emarginatus, Bethalus
319, 320, 322, 330
. 124
319
323, 325. 326, 333
. 204
223-227, 245
. 224
. 199
. 229
. 214
59-64, i PL
. 95-102, i PL
225
225
232
232
225
250, 253, 270
. 220
. 209
. 189-2 4 6
194
212, 242
. 222
emarginatus, Scotophilus .
Emballonuridae . 317,
322, 323,
328, 329,
Emballonuroidea
Emoa ....
Emoia ....
emunitus, Armadillo
emunitus, Bethalus .
Ennurensis
episimus, Hekelus .
epops, Upupa
Eptesicus
erinaceus, Armadillo
erinaceus, Spherillo .
erraticus, Cyamus .
erythroleucus, Armadillo .
Eschrichtius .
esterelanum, Armadillidium
esterelanus, Cylisticus
Ethelum
Eubalaena
Eubelidae . . 191
Eubelum
Euglypha
Euglyphidae .
Euplotes . 95-102, i
Euporcellio
eurystomus, Euplotes 118,
evansi, Porcellio
exarata, Campylaspis 250,
Exzaes .
173
318, 319, 320, 321,
324. 325, 326, 327,
330. 332, 333, 334
329, 334
. 56
53-58
. 222
. 222
. . . 193
213
37. 42
74. 134
. 229
. 229
. 67-68, 71
. 220
70. 71
. 2O9
. 192
213-214
67, 68, 69, 71
, 213-218, 242-244
214, 215, 242, 243
103-112, 2 Pis.
. IO3-II2, 2 Pis.
PL; 113-126, i PL
. 207
119, I2O, 121, 124
. 199
254, 266, 284-285
. 212
falabae, Scotoecus . . 169, 178, 183, 184
falabae, Scotoecus hindei . . . 183-184
Falculea. . . . . . -42
famosa, Nectarinia . . . .42
feae, Armadillo . . . . .228
feae, Pericephalus ..... 228
festivus, Armadillo ..... 225
festivus, Diploexochus .... 225
ficorum, Porcellio . . . . .199
ficulneus, Porcellio . . . . . 199
finmarchica, Cythereis . . . 83, 84
finmarchica, Finmarchinella . 83, 84-85,
86, Pis. 1-2
finmarchica, Hemicythere . . .84
Finmarchinella . . . 83-94, 2 Pis.
fissifrons, Pagana . . . . .206
flavocinctus, Porcellio . . . . 199
flavoscutatum, Armadillidium . . . 209
flavovittata, Porcellio . . . . 199
flavus, Armadillo ..... 220
fongosiensis, Anchicuboris . . .219
fontium, Protracheoniscus . . . 206
formosana, Nagara .... 205
formosanus, Nagurus .... 205
fragilis, Agnara . . . . .205
frascatensis, Orthometopon dalmatinus . 206
INDEX
frigida, Campylaspis . . . 250, 255
fritschei, Reductoniscus .... 229
frontalis, Sphaeroniscus . . . .218
frontalis, Spherillo . . . . .230
frontexcavatum, Armadillidium . . 209
frontirostre, Armadillidium . . . 209
frontosus, Buddelundia .... 223
frontosus, Metoponorthus . . . 196
furcatum, Armadillidium .... 209
furcatus, Diploexochus . . . .225
Furipteridae . 318, 319, 320, 326, 327, 328, 330
fuscomarmoratus, Metoponorthus . .196
fusiformis, Campylaspis . . . 250, 252
grisescens, Spherillo.
grossus, Armadillo .
grossus, Spherillo
guttata, Campylaspis
Gymnastinops
230
230
. 230
250, 251, 258-259
39
galapagoensis, Cubans .
galapagoensis, Diploexochus
galeatus, Armadillo .
galeatus, Pericephalus .
gallicus, Porcellio . .
Gelsana . . . .
gemmulatus, Porcellio .
Gerufa ....
Gerutha ...
gestroi, Armadillidium .
gibbosus, Eschrichtius .
gigas, Armadillidium simoni
gigas, Cubaris ..
gigas, Diploexochus .
glabra, Campylaspis .
glacialis, Eubolaena. .
glacialis, Eubolaena glacialis
glacialis australis, Eubalaena
glacialis glacialis, Eubalaena
glacialis japonica, Eubalaena
glebulosa, Campylaspis .
Globicephala ...
globosa, Campylaspis
globus, Synarmadillo .
Glomerulus ...
glomus, Armadillo . .
glomus, Diploexochus .
Glossophaga . . .
gordoniensis, Diploexochus
gracilior, Merulana translucida
gracilipennis, Cylisticus .
gracilis, Cyamus . .
grandinatum, Armadillidium
graniger, Porcellio . .
granulata, Campylaspis .
granulatum, Armadillidium . .
granulatum peloponnesiaca, Armadillidium
granuliferus, Porcellio ...
gravei, Metoponorthus . . .
... 225
. . . 225
... 228
... 228
. . . 199
. . .214
... 199
... 194
.. 214-215
. . . 209
. . 70, 71
. . .211
... 225
. . .225
. 250, 252, 261
. . .69
. 68, 69, 71
. 67, 68, 69, 71
. 68, 69, 71
. . 69, 71
. 250, 255, 293-295
... 70, 71
250, 255, 291, 295, 297
. . . .231
.... 228
. . . .225
.... 225
. . . -325
. 225
. 228
.192
68, 71
. 209
. 199
250, 252
.210
209
200
196
Greenwood, P. H
grenadensis, Armadillo
grenadensis, Venezillo
greyii, Scoteinus .
griseoalbus, Armadillo
griseoalbus, Bethalus
139-i65
. 232
.232
. 173
.222
.222
Haloporcellio ..
Haplarmadillo . .
Haplochromis ...
harpa, Euplotes . .
harsadiensis, Armadillo .
hartae, Campylaspis .
hebridarum, Spherillo .
Hedley, R. H. . .
Hekelus . . .
Hemichromis . .
Hemicythere ...
"Hemicytherinae" . .
Hemignathus . . . .
Hemilepistus ....
Hemiporcellio . . .
hermagorensis, Protracheoniscus
Hermanites . . . .
herzegovinensis, Porcellio . .
herzegovinensis, Porcellium .
Heteralocha ....
Hiallum . . . .
hilgendorfi, Eubelum . .
hilgendorfi, Hiallum . .
... 194
. . -215
.. 139-I&5
. . 101, 119
... 220
. . 250, 253
... 230
103-1 12, 2 Pis.
. . .213
. . .141
84, 86, 87, 88, 89
. . .88
. . 5, 42
.. 194, 237
. . .195
206
86
206
206
i PI.
215
215
215
1-
Hill, J. E. 73-82; 127-138; 167-188; 301-336
hindei, Nycticeius hirundo . . .182
hindei, Scotoecus . 169, 171, 172, 175, 178,
179-i8o, 181, 182, 183, 184, 185, 186
hindei, Scotoecus hindei 179, 180-i82, 183, 184
hindei, Scotoecus hirundo . 181, 182, 183
hindei albigula, Scotoecus 180, 181, 182-i83
hindei artinii, Scotoecus . . . .182
hindei falabae, Scotoecus . . . 183-1 84
hindei hindei, Scotoecus 179, 180-i82, 183, 184
Hipposideridae 317, 318, 319, 320, 322, 323,
325, 326, 327, 328, 329, 330, 332, 333
Hipposideros ...... 303
hirsutulus, Protracheoniscus . . . 206
hirticornis, Gerufa . . . . . 194
hirtum, Armadillidium .... 209
hirtus, Agabiformius . . . . 193
hirtus, Lucasius ..... 193
hirundo, Scotoecus . . 169, 171, 172, 175,
177-179, 181, 183, 184, 185, 186
hirundo, Scotophilus . . . . 177
hirundo hindei, Nycticeius . . .182
hirundo hindei, Scotoecus . 181, 182, 183
hispida, Metoponorthus . . . .196
hispida, Porcellio . . . . .196
hispida, Porcellionides . . . 196
hispidus, Ennurensis . . . . 193
holosericeus, Amblyrhamphus . . -37
horrida, Campylaspis . . 250, 255, 292
342
INDEX
horridoides, Campylaspis .
horsfieldii, Myotis .
horsfieldii, Myotis horsfieldii
horsfieldii horsfieldii, Myotis
horsfieldii lepidus, Myotis.
hughscotti, Periscyphis
Huia ....
Hurley, D. E.
hybridum, Armadillidium
hybridum, Schizidium
Hyperoodon .
hypotoreus, Spherillo
Hypotrichida .
hypselos, Diploexochus
hypsinephes, Diploexochus
250, 255, 292-293
. 132
. 132, 137
. 132, 137
132
. 216
1-48, i PI.
59-64, i PI. ; 65-72
. 213
. 213
. 71
230
95-102, i PI.; 113
. 225
. 225
Ignamba . . . . . .215
ignavum, Eubelum . . . . .214
imbutus, Porcellio ..... 200
imbutus pellegrinensis, Porcellio . . 200
immotus, Armadillo .... 220
immsi, Hemiporcellio . . . . 195
immsi, Porcellio . . . . .195
inaequalis, Buddelundia .... 223
incanus, Porcellio ..... 200
Inchanga ...... 195
incicus, Scleropactes . . . .218
incisa, Nagara ..... 205
incisus, Nagurus ..... 205
inconspicuus, Armadillo .... 220
indentatus, Euplotes .... 101
ingens, Spherillo ..... 230
iniqua, Merulana ..... 228
iniquus, Spherillo ..... 228
inkystans, Euplotes . . . . 101
inornata, Campylaspis . . . 250, 251
instrenuum, Eubelum . . . .214
integer, Armadillo .... 220
intercalarius, Porcellio .... 200
intermedia, Campylaspis 250, 255, 277, 289, 297
intermixtus, Armadillo .... 220
interpolator, Porcellio .... 200
Isocyamus ..... 59, 70, 71
japonica, Barentsovia . . . 91-92
japonica, Eubalaena glacialis . . 69, 71
japonica, Finmarchinella . . . 91-92
japonica, Nereina . . . . 83, 91
jelskii, Porcellio ..... 200
johnstoni, Campylaspis . . . 250, 253
johorensis, Tadarida . . . 136-137
Jones, N. S 247-30O
jonesi, Campylaspis . . . 250, 253, 263
kaokoensis, Diploexochus . . . 225
kiiensis, Campylaspis . . . 250, 252
Kerivoula
Kerivoulinae .
Kisuma .
kogmani, Diploexochus
Krafft, J. I. .
kuhlii, Scotophilus .
kunenensis, Periscyphis
kunenensis, Periscyphops
. 328
319, 320, 325
232
225
103-112, 2 PIS.
174
. 216
. 216
labiatus, Buddelundia .... 223
lacteolus, Metoponorthus .... 196
Laephotis ..... 73-82
laevigata, Campylaspis 250, 251, 258, 260-26i
laevigatus, Buddelundia .... 223
laevis, Gerutha . . . . .214
laevis, Porcellio . . . . .201
Lagenorhynchus .... 59, 71
lamellatus, Porcellio . . 191, 194, 201
laminigerum, Armadillidium . . .210
larii, Trachelipus ..... 207
larii, Tracheoniscus ..... 207
larii aternanus, Trachelipus . . . 207
larii aternanus, Tracheoniscus . . . 207
Lasiurus ...... 328
lateralis, Buddelundia .... 223
laticarpa, Campylaspis . . . 250, 254
laticauda, Mahehia .... 195
laticauda, Tura ..... 204
latidactyla, Campylaspis . . . 250, 253
Latidens . . . . . -303
latifrons, Armadillo . . . .222
latifrons, Bethalus . . . . .222
latissimus, Periscyphis . . . .216
latissimus, Porcellio. .... 200
latus, Euplotes patella .... 124
latus, Protracheoniscus .... 206
legendrei, Campylaspis . . . 250, 253
lekaguli, Hipposideros .... 303
lentus, Spherillo ..... 230
lepidus, Myotis . . . . .132
lepidus, Myotis horsfieldii. . . .132
lepidus group, Rhinolophus . . .129
lepidus subgroup, Rhinolophus . .129
lepineyi, Porcellio ..... 200
Leptotrichus . . . . . 195, 237
leucocephalus, Microcercus . . -215
leucocephalus, Periscyphis . . -215
leucorhamphus, Cacicus . . . -37
leucostictus, Sarotherodon . . .163
lifuensis, Armadillo ..... 220
lifuensis, Cubaris . . . . .220
limbata, Periscyphis . . . .216
limbatus, Bethalus . . . .222
limenites, Diploexochus . . . .225
Lincoln, R. J.. 59-64, I p l-.' 66-72; 189-246
lineare, Trinema . . . . -103
Liponycteris . . . . . -323
littoralis, Metoponorthus . . . .196
INDEX
343
loati, Haplochromis . . . . .162
Lonchorhina ...... 326
longicauda, Porcellio .... 200
longipennis, Porcellio .... 200
longipes, Armadillo .... 225
longipes, Diploexochus .... 225
loyaltiensis, Emoia samoensis . . 53-58
lubricum, Eubelum . . . . .214
Lucasius . 191, 193, 195, 197, 201, 237
lugubris orarum, Porcellio . . .201
lugubris vizzavonensis, Porcellio . .201
lusitanus, Porcellio .... 202
lusitanus, Porcellio scaber . . . 202
lutshniki, Trachelipus .... 207
lutshniki, Tracheoniscus .... 207
Lygosoma . . . . . -56
Lyprobius ...... 205
macconneli, Haplochromis . 141, 149, 153,
154-i6i, 162, 164
macmahoni, Armadillo .... 220
macrocneme, Miniopterus medius . 135, 137
macrodens, Bethalus .... 222
macrophthalma, Campylaspis . 250, 253, 264
macrops, Gerufa ..... 194
maculata, Balloniscus .... 205
maculata, Campylaspis . . . 250, 255
maculata, Mahehia . . . . 195
maculata, Philoscia .... 205
maculatum cingendum, Armadillidium . 208
maculipes, Porcellio. .... 200
maculosa, Pagana ..... 206
maculosus, Spherillo .... 230
madagascariensis, Agnara . . . 205
madagascariensis, Metoponorthus . . 205
magna, Sturnella . . . . -37
magnicirratus, Euplotes . . . . 101
magnificus, Porcellio .... 200
mahagiensis, Haplochromis . . .162
Mahehia. ...... 195
makuae, Diploexochus .... 226
Mammalia . 127-138; 167-188; 301-336
mammillata, Sunniva . . . -213
mansa, Campylaspis 250, 254, 283, 286-287
marcidus, Armadillo . . . .228
marcidus, Pericephalus .... 228
mareoticum, Armadillidium . . .210
marginalis, Uramba. .... 205
marginatus, Spherillo . . . -230
marginenotatus, Porcellio. . . . 200
marginepilosa, Gerutha . . . -215
marinensium, Armadillidium . . .210
marmorata, Gerufa . . . . .194
marmoratus, Synarmadillo . . .231
marquesarum, Spherillo .... 230
marquesarum, Xestodillo .... 230
marshalli, Rhinolophus .... 303
mauritanica, Campylaspis . 250, 251, 261
mayeti, Armadillo . . . . .220
medius, Miniopterus . . . -135
medius macrocneme, Miniopterus . 135, 137
Medway, Lord .... 53-58
Megadermatidae . . 318, 319, 320, 322,
325, 326, 327, 329, 330, 332
Megaptera . . . .66, 67, 70, 71
mehelyi, Protracheoniscus . . . 206
mehelyi, Protracheoniscus politus . . 206
meiringi, Diploexochus . . . .226
melaena, Globicephala . . . 70, 71
melanocephalus, Chironax . 127-129, 137
melanurus, Acaeroplastes . . -193
melanurus, Metoponorthus . . . 193
meleagris, Metoponorthus . . . 197
meleagris, Metoponorthus pruinosus . 197
Merulana ...... 228
Mesarmadillo . . 213, 214, 215, 243
Metoponorthus . . 193, 196-197, 199,
200, 204, 205, 206, 237, 238
197
215, 243
318, 322, 324, 325
325
228
. 228
. 215
. 201
319, 327
135. 137. 328
250, 253
. 129, 131
. 213
. 129
193
. IOI
213
193
. 22O
. 2I 4
. 214
113, 114, 115-117,
Il8, 119, I2O, 121, 123
. 201
moebiusi quadricirratus, Euplotes . 113, 123
mohamedanicum, Armadillidium . .210
Molossidae . . 318, 319, 320, 322, 324,
325, 326, 327, 328, 329, 330
monocellatus, Haplarmadillo . . -215
monoceros, Monodon . . 68, 69, 71
Monodon .... 68, 69, 71
monodontis, Cyamus . . . 68, 71
montagui, Diploexochus .... 226
montana, Gerufa . . . . . 194
montanus, Armadillo .... 220
montanus, Porcellio. .... 203
montanus, Porcellio spinipennis . . 203
monticola, Buddelundia .... 223
montivagus, Armadillo .... 230
montivagus, Spherillo .... 230
Mica
Microcercus
Microchiroptera
Micronycteris .
microps, Armadillo .
microps, Glomerulus
microps, Ignamba .
mildei, Porcellio
Miniopterinae .
Miniopterus
minor, Campylaspis
minor, Rhinolophus.
minor, Sunniva
minor subgroup, Rhinolophus
minuta, Angara
minuta, Euplotes
minuta, Pareluma .
minutus, Agabiformius
miser, Armadillo
modestum, Ethelum
modestus, Mesarmadillo .
moebiusi, Euplotes
moebiusi, Porcellio
344
INDEX
Mormoopidae .
319. 320, 323. 325. 326.
327, 328, 329, 330, 332, 333
325
. 222
250, 256, 297-2QQ
. 2IO
. 320
205
. 205
65, 71
100, 117
132, 137. 303
Mormoops
tnucidus, Bethalus .
multinodosa, Campylaspis
muricatum, Armadillidium
Murininae
mus, Lyprobius
mus, Uramba
musculus, Balaenoptera .
mutabilis, Euplotes
Myotis ....
myrmecophilus, Metoponorthus myrmecophilus
196
myrmecophilus, Myrmeconiscus myrmecophilus
196
myrmecophilus myrmecophilus, Metoponorthus
196
myrmecophilus myrmecophilus, Myrmeconiscus
196
myrmicidarum, Metoponorthus . .196
Mystacinidae . ' .
mystica, Sunniva
Mysticeti
mysticetus, Balaena
Myzopodidae .
318, 319, 320, 324,
325. 327. 329, 33
. 213
. 71
67, 71
318, 320, 326, 327, 330
Nagara .
Nagurus .
namibensis, Laephotis
nana, Nagara .
Nannastacidae
nanus, Diploexochus
nanus, Nagurus
narentanus, Porcellio
nasatum, Armadillidium
. 205
. 205, 239
73, 75-76, 77, 79, 80
. 205
. 249
. 226
205
. 201
. 212
nasatum sorrentinum, Armadillidium . 210
nasatus, Cylisticus ..... 192
natalensis, Inchanga .... 195
Natalidae . 318, 320, 326, 327, 328, 330
naupliensis, Porcellio
Neale, J. W. .
nebulosus, Diploexochus
Nectarinia
Neocyamus
Nereina .
Nesodillo .
nicklesi, Porcellio
nicobaricus, Armadillo
nicobaricus, Spherillo .
nigra, Emoia .
nigra, Porcellio
nigricans, Periscyphis
nigripes, Buddelundia .
nigrobrunneus, Metoponorthus
nigromarginatus, Armadillo
nigropunctatus, Periscyphis
nigropunctatus, Synarmadillo
. 2OI
83-94, 2 Pis
. 226
. 42
59, 70, 71
83, 86, 87, 88, 90, 91
. 228
f . . 201
230
230
53
. 201
. 217
. 223
. 196
. 221
231
. 231
niloticus, Sarotherodon
nitens, Campylaspis.
nitidulus, Armadillidium .
nobilis, Spherillo . .
Noctilio .
Noctilionidae . . 318,
325, 326
noctulinus, Nycticejus
noctulinus, Vespertilio
nodosus, Cyamus
nodosus, Hemilepistus
nodulosa, Campylaspis
normani, Lucasius .
normani, Porcellio .
Normanicythere
novaeangliae, Megaptera .
nuda, Campylaspis .
nuda, Cythere clathrata .
Nyctalus
Nycteridae . 318, 319,
325. 326, 327
Nycticeius
Nyctophilinae
140, 149
250, 251, 256, 258
. 210
230
329
319, 320, 323, 324,
327, 329, 330, 332
i?4
. 172, 174
68-69, 7i
194
. 250, 255
. 2OI
. . 201
8 5
66, 67, 70, 71
250, 251, 256-258
86, 87, 88
. 171
320, 322, 323, 324,
329, 330, 332, 333
169, 170, 171, 173,
174, 175, 182, 186
317, 322
obliquidens, Diploexochus . . . 226
obliquipes, Spherillo . . . -230
obsoletus, Porcellio ..... 201
obtusa, Angara . . . . . 193
obtusifrons, Porcellio . . . .201
obtusiserra, Porcellio .... 201
obtusus, Agabiformius .... 193
occidentals, Protracheoniscus . . .196
ocellatus, Porcellio . . . . .201
octocarinatus, Euplotes . . . .124
octocirratus, Euplotes . . . 100, 101
odherni, Armadillidium . . . .210
odherni, Armadillidium peraccai . .210
Odontoceti . . . . . -71
oertzeni, Armadillidium . . . .213
oertzeni, Schizidium . . . .213
officinalis syriaca, Armadillo . . .221
Ogden, C. G 103-1 12, 2 Pis.
oliveti, Armadillidium . . . .210
olivieri, Porcellio . . . . .198
Oniscoidea ..... 189-246
opacus, Buddelundia .... 223
opihensis, Actoecia . . . .218
orarum, Porcellio lugubris . . . 201
orarum, Porcellio orarum . . .201
orarum orarum, Porcellio . . .201
orarum vizzavonensis, Porcellio . . 201
orientalis, Campylaspis . . . 250, 251
ormeanum, Armadillidium . . .210
ornatus, Scotomanes . . . . 173
Orodillo 228
orphanus, Diploexochus .... 226
Orthometopon ..... 206
Ostracoda .... 83-94, 2 Pis.
INDEX
345
Ourachaerus .
ovalis, Campylaspis .
ovalis, Cyamus
ovampoensis, Cubaris
ovampoensis, Diploexochus
oxyzomus, Armadillo
oxyzomus, Cubaris
232
250, 253
69, 71
. 226
. 226
. 221
. 221
pachytos, Diploexochus
pacifica, Campylaspis
pacificus, Cyamus
paeneglabra, Campylaspis
Pagana .
palaestinus, Hemilepistus
pallasi, Armadillidium
palliata, Falculea
pallidus, Cylisticus .
pallidus, Lucasius
pallidus, Porcellio
pallidus, Scotoecus .
pallidus, Scotophilus
Pandanus
panurus, Armadillo .
panurus, Bethalus .
papillata, Campylaspis
papillosa, Kisuma
Paracubaris
paradoxolophus, Rhinolophus
Paraperiscyphis
Pareluma
parietinus, Porcellio
parkei, Euplotes
parvus, Armadillo
parvus, Porcellio
parvus, Scutocyamus .
parvus, Spherillo
patella, Euplotes
patella latus, Euplotes
paucinodosa, Campylaspis
paucispina, Campylaspis .
pauper, Porcellio
pauperculus, Diploexochus
pellegrinense, Armadillidium
pellegrinensis, Porcellio imbutus . . 200
Pelmatochromis . . . . .141
peloponnesiaca, Armadillidium granulatum 209
peltatus, Spherillo . . . . -230
penicilliger, Haloporcellio . . . -194
peraccae, Armadillidium . . . .210
peraccai, odherni, Armadillidium . .210
Pericephalus ...... 228
Periscyphis . . 215,216-217,218,231,243
Periscyphops . . .216,217-218,243,244
perkinsi, Armadillo . . . . .231
perkinsi, Spherillo . . . . .231
petraeum, Armadillidium . . . .210
petronius, Haplochromis . . . 161, 163
phaeacorum, Porcellio rathkei . . . 207
. 226
. 250, 251
70, 71
250, 251, 258, 261
. 206
194
. 2IO
. 4 2
. 192
. 195
195
171, 172-174, 175, 186
169, 170, 172, 173
195
. 222
. 222
250, 255, 296
232
. 218
303
. 216
213
. 2OI
119-121, 122, 123, 124, PI. I
. 230
. 201
59-64, i PL; 71
230
97. 99. IO . 117. 124
. 124
2 5. 255, 291-292
250, 254, 275-277
. 201
. 226
. 210
phaeacorum, Trachelipus .
Phalaba ....
phaleronensis, Metoponorthus .
phaleronensis, Orthometopon
phaleronensis, Porcellio
pharyngalis, Haplochromis
Philetor
Philoscia ....
philoscoides, Metoponorthus
Phoeniculus ....
Phoniscus ....
Phyllostomatidae 317, 319, 320,
327.
Phyllostomatoidea .
physalus, Balaenoptera
Physeter ....
physeteris, Neocyamus
pictus, Rotungus
piger, Armadillo
pila, Gerutha ....
pileus, Campylaspis .
pilosa, Campylaspis . 250,
pilosa, Gerutha
pilularis, Armadillo .
pilum, Eubelum
Pipistrellus . . . 132-
Pisces .....
pisum, Armadillo
pisum, Diploexochus
planarius, Porcellio .
Plataoniscus ....
Platycyamus ....
platyuropus, Campylaspis .
plicata, Campylaspis 250, 254,
plicata, Tadarida
plumbeus, Cylisticus
plumbeus bergomatius, Cylisticus
plumbeus umbricus, Cylisticus .
plumipes, Euplotes .
politus, Leptotrichus
politus mehelyi, Protracheoniscus
pollex, Diploexochus
Polyacanthus ....
polythele, Diploexochus .
pontremolensis, Cylisticus .
porcata, Campylaspis . 250, 252,
Porcellidium ....
Porcellio 191, 194, 195, 196,
. 207
206, 239, 240
. 206
. 206
. 206
161, 163
134-135. 137
205
. 196
37-45
135-136, 137
322, 325, 326,
329, 330. 332
329
. 65, 66, 71
66, 67, 70, 71
70, 71
. 218
. 221
. 215
250, 252
254, 271-272
215
. 221
. 215
134. 137. 303
. 139-165
. 226
. 226
. 202
. 205
59.70-71
250, 252, 259
280-282, 289
136, 137
. 192
. 192
. 192
117, 119, 124
195
. 206
. 226
. 229
. 226
Porcellionidae
Porcellionides .
Porcellium .
porphyrivagus, Caeroplastes
porphyrivagus, Metoponorthus .
portofinense, Armadillidium
praeustus, Porcellio .
pretoriensis, Armadillo
pretoriensis, Bethalus
Prionoplus .
procerus, Hemignathus
. 192
261-2&3, 265
. 206
197-204, 206,
207, 238, 239
I9L 193-205, 237-239
196, 204
. 206
193
193
210
2O2
222
222
4
42
346
INDEX
Protozoa 95-102, i PI.; 103-112, 2 Pis.;
113-126, i PI.
Protracheoniscus . . . 196, 206-207
provincialis, Porcellio .... 202
proximatus, Armadillo . . . .221
pruinosus, Metoponorthus . 199, 200, 204
pruinosus anconanus, Metoponorthus . . 197
pruinosus meleagris, Metoponorthus . . 197
Psarocolius ..... 37, 39
pseudoratzeburgi, Euporcellio . . . 207
pseudoratzeburgi, Porcellio . . . 207
pseudoratzeburgi, Trachelipus . . . 207
pseudorcae, Syncyamus . . . .62
Pseudotracheata .... 189-246
Pteropodidae . . . . . -325
pujetanum, Armadillidium . . .211
pujetanus, Porcellio .... 202
pulchella, Campylaspis . . . 250, 251
pulcher, Periscyphis . . . ..217
pumila, Campylaspis . . 250, 252, 263
pumilus, Armadillo ..... 232
pumilus, Rhinolophus cornutus . . -131
pumilus, Venezillo ..... 232
purpurascens, Eluma . . . .212
purpurascens, Spherillo . . . .231
purpureus, Phoeniculus . . . -37
purpureus, Porcellio ..... 202
pusillus, Rhinolophus . . 129-131, 137
pusillus group, Rhinolophus . . 129,131
pustulosa, Campylaspis . . . 250, 255
pygmaeus, Armadillo . . . .221
pygmaeus, Periscyphis . . . -231
pygmaeus, Synarmadillo . . . -231
pyrenaeus, Porcellio ..... 202
Pyrgoniscus ...... 232
pyriformis, Tetrahymena . . . -97
Pyrrhocorax . . . . . -37
pyrrhocorax, Pyrrhocorax . . -37
quadricirratus, Euplotes moebiusi . 113,123
quadrimaculatus, Diploexochus . . .226
quadrimaculatus, Mesarmadillo . -215
quadrimaculatus, Periscyphis . . .217
quadriplicata, Campylaspis . . 250, 254
quadriserriatum, Armadillidium . .211
quadritracheata, Buddelundia . . . 223
quietum, Eubelum . . . . .214
quinquecarinatus, Euplotes . . 97, 101
quinquepustulatum, Armadillidium . .211
ragusae, Porcellio ..... 202
raikovi, Euplotes . . .99, 100, 101, 123
rariseta, Euplotes . 95-IO2, i PL; 115, 118
rathkei phaeacorum, Porcellio . . . 207
razae, Alauda .
reaumuri, Hemilepistus
redacta, Campylaspis . 250,
Reductoniscus
reflexum, Ethelum .
reflexus, Mesarmadillo
rehobotense, Armadillidium
Reptilia ....
reticularis, Prionoplus
reticulata, Campylaspis
Rhinolophidae 318,319,320,
327. 328,
Rhinolophoidea
Rhinolophus .
Rhinopoma . . 304, 316,
Rhinopomastus
Rhinopomatidae3i6, 317, 318,
323, 324, 325,
rhodesiae, Bethalus .
rhodesiensis, Diploexochus
ribauti, Sphaerobathytropa
riparium, Armadillidium .
robusta, Arachnothera
rohui, Philetar
rohui, Philetar brachypterus
rosai, Armadillidium
roscida, Campylaspis
roscoffensis, Euplotes
rostellata, Campylaspis 250
rostrata, Campylaspis
rotunda, Euglypha .
Rotungus
Rousettus
rubicunda, Campylaspis . 250
rubromaculata, Campylaspis,
rudolfianus, Haplochromis
153,
rufa, Campylaspis
rufescens, Diploexochus
ruficauda, Periscyphis
rufobrunneus, Agabiformius
rufomarginatus, Spherillo .
rugifrons, Buddelundia
rugulosus, Cubaris .
rugulosus, Spherillo .
rupta, Campylospis .
sabuleti, Metoponorthus
sabulifer, Porcellio .
Saccolaimus .
Saccopteryx .
sagamiensis, Campylaspis
Saidjahus
saldanhae, Diploexochus
salimalii, Latidens .
. 41
194
254, 274-275, 278
. 229
. 214
. 214
. 211
53-58
4
. 250, 253
323, 324, 325, 326,
329, 33, 332, 333
332
129-132, 137, 303
,321,323,332,333
. 38,42,45
319, 320, 321, 322,
326, 328, 329, 330,
332, 333. 334
. 222
. 227
. 218
. 211
. 4 2
134. 135
135
. 211
. 250, 254
. IOI
, 253, 267-268, 270
250, 255, 268, 289
103, 108
. 218
. 129
,251, 259-26o, 261
250, 255
141, 142-150, 152,
155. 156, 157. 158.
159, 160, 161, 162
. 250, 252
. 227
. 217
193
. 231
. 223
. 231
231
. 250, 252
. 197
. 202
323
323
250, 255
. 229
. 227
303
INDEX
347
salisburyensis, Diploexochus . . .227
saltuum, Porcellio ..... 202
samoense, Emoa . . . . .56
samoense, Lygosoma . . . -56
samoensis, Emoia .... 54, 55, 56
samoensis loyaltiensis, Emoia . . 53-58
sanctum, Armadillidium . . . .211
sanfordi, Emoia .... 55, 56
sarajevensis, Porcellio .... 202
sarasini, Nesodillo . . . . .228
sarculatus, Porcellio ..... 202
Sarotherodon .... 148, 149, 163
sauteri, Orodillo . . . . .228
savonense, Armadillidium . . . .211
saxicola, Haplochromis . . . -157
saxonicus, Protracheoniscus . . . 207
saxonicus carpathicus, Protracheoniscus . 207
scaber, Porcellio .... 199, 201
scaber lusitanus, Porcellio . . . 202
scaberrimum, Armadillidium . . .211
scabrum, Armadillidium . . . .211
scammoni, Cyamus .... 70, 7 1
schellenbergi, Nesodillo .... 228
schirasi, Hemilepistus .... 194
Schizidium . . . . . -213
schlieffenii, Nycticeius . . . . 175
schlieffenii, Scoteinus . . 173, 1 8 1, 182
Scincidae ..... 53-58
Scleropactes . . . . .218, 244
Scoteinus . . 170, 171, 173, 181, 182, 186
Scotoecus ..... 167-i88
Scotomanes . . . . . . 173
Scotophilus . . . 169, 170, 171, 172,
173, 174, 175, 177
Scotozous . . . . . .169
Scutocyamus . . 59-64, l Pl- 7 1
serratipes, Campylaspis . . . 250, 254
serratum, Armadillidium . . . .211
setaceus, Spherillo . . . . .231
sexfasciatus, Metoponorthus . . . 197
sexfasciatus asifensis, Metoponorthus . . 197
sharpi, Armadillo . . . . .221
siculorum, Armadillidium badium . . 208
silvanus, Periscyphops . . . .217
silvarum, Armadillo .... 227
silvarum, Diploexochus . . . .227
similis, Campylaspis . . . . 250,252
simoni, Armadillidium . . . .211
simoni gigas, Armadillidium . . .211
simplex, Synarmadillo .... 232
simulator, Porcellio ..... 202
sinuosa, Campylaspis . . . 250, 253
Skink . .... 53-58
sociablis, Porcellio ..... 203
societatis, Pipistrellus . . . 133, 134
solitarius, Cacicus .... 36, 37
sollers, Spherillo . . . . .231
sordidus, Porcellio ..... 203
sorrentinum, Armadillidium nasatum . 210
spatulata, Porcellio . . . . .203
spatulatus, Porcellio .... 200
speciosa, Campylaspis . . . 250, 253
speciosa, Campylaspis costata . . .266
speiseri, Emoia .... 54, 55
Sperillo ....... 245
speyeri, Armadillidium . . . .212
Sphaerobathytropa . . . . .218
Sphaeroniscidae . . . 191, 218, 244
Sphaeroniscus . .... 218,244
Spherillo . . .220,228,229-231,245
sphinx, Haloporcellio . . . 191, 194
spicatus, Spherillo . . . . .231
spinicornis, Porcellio .... 203
spinifera, Campy laspides . . . .270
spiniger, Acanthoniscus . . . .219
spinipennis, Porcellio .... 203
spinipennis, Porcellio spinipennis . 202, 203
spinipennis montanus, Porcellio . . . 203
spinipennis spinipennis, Porcellio . 202, 203
spinipes, Porcellio ..... 203
spinosa, Akermania . . . . .219
spinosa, Campylaspis . 250, 253, 267, 270
spinosus, Circoniscus . . . .218
spinosus, Paracubaris . . . .218
spretus, Porcellio ..... 203
squamatus, Periscyphops . . . .217
squamifera, Campylaspis . 250, 255, 293, 295, 296
squamosus, Periscyphops . . . .217
stebbingi, Paraperiscyphis . . .216
steenbrasi, Diploexochus . . . .227
Steno . . . . . . 70, 71
Stenoderminae . . . . .319
sticta, Campylaspis . . 250,253,264-265
stipulatum, Eubelum . . . .214
stolikanum, Armadillidium . . .212
straeleni, Haplochromis . . . .162
strelkovi, Euplotes ..... 101
striata, Campylaspis . . . 250, 252
stricticauda, Armadillo .... 222
stricticauda, Bethalus . . . .222
strigosa, Euglypha . . . 103-1 12, PI. 2
Sturnella . . . . . -37
Sturnus ....... 37
subbadius subgroup, Rhinolophus . .129
subdentatum, Armadillidium . . .212
suberorum, Cylisticus .... 192
subjaponica, Cushmanidea . . .86
submersa, Campylaspis . 250, 255, 290-291
subterraneus, Desertoniscus . . . 193
subtransversus, Periscyphis . . .217
succinctus, Porcellio .... 203
sulcata, Campylaspis . . 250, 253, 275
sulcatus, Buddelundia .... 223
sumatranus, Rhinolophus acuminatus . 132
Sunniva ...... 213
sylvatica, Exzaes . . . . .212
Synarmadillo ..... 231-232
Syncyamus ..... 59, 62
syriaca, Armadillo omcinalis . . .221
szechwanus, Rhinolophus blythi . -131
INDEX
tabularis, Diploexochus . . . .227
Tadarida . . . 136-137
tamei, Periscyphis . . . . .217
Taphozous . . 323, 326, 328, 333
tarangensis, Spherillo . . . .231
tardus, Mica ...... 197
tardus, Porcellio ..... 197
taschkententis, Protracheoniscus . . 207
tegulatus, Euplotes .... 101, 124
temminckii, Scotophilus . . . . 174
tendanum, Armadillidium . . .212
tenuipunctatus, Armadillo . . .221
Testacea .... 103-112, 2 Pis.
testacea, Tura ..... 204
Tetrahymena . . . . . -97
thetidis, Campylaspis . . . 250, 255
thompsoni, Campylaspis . . . 250, 252
thompsoni, Platycyamus . . . 70-7 1
thomsoni, Cyamus . . . . .70
thonglongyai, Craseonycteris . 301-336
Thyropteridae . 318,319,320,326,327,330
Tura ....
turkanae, Haplochromis .
. 204
141, 149, 150-153,
155. i 60
tigris, Armadillidium
Tilapia .
tirolense, Armadillidium
tomentosus, Buddelundia
Toradjia
torulosa, Campylaspis
tosaensis, Hermanites
trachealis, Porcellio .
Trachelipidae .
Trachelipus ....
Tracheoniscus
tradouwi, Bethalus .
translucida, Merulana
translucida gracilior, Merulana .
translucidus. Armadillo
transmutatus, Porcellio
transsilvaticus, Cylisticus .
transvaalensis, Polyacanthus
travancoria, Nagara .
travancorius, Nagurus
Trematocranus
triangulifera, Uramba
triarticulatus, Periscyphis .
triarticulatus, Periscyphops
tricirratus, Euplotes affinis
trifolium, Armadillo.
trilobatus, Porcellio .
Trinema. ....
triplicata, Campylaspis
trisulcatus, Euplotes
trivialis, Periscyphis
trochilirostris, Campylorhamphus
truncorum, Armadillo
truncorum, Venezillo
. 212
148, 149
. 212
223
. 242
250, 255, 295-296
203
191, 205-207, 239-240
. 207
. 207
. 222
. 228
. 228
. 228
203
. 192
. 229
205
205
. 160
205
. 218
. 218
113, 119, 123
. 221
2O3
. 103
250, 253
. IOI
. 217
. 42
232
232
tuberculatus, Mesarmadillo
tubulata, Campylaspis
tuffraui, Euplotes
tugelae, Diploexochus
tunetanum, Armadillidium
215
250, 252
101, 124
. 227
. 212
umbensis, Campylaspis . . . 250, 253
umbricus, Cylisticus plumbeus . . .192
undata, Campylaspis .250, 253, 283, 287, 289
undulata, Periscyphis . . . .217
uniformis, Sunniva . . . . .213
uniplicata, Campylaspis . . . 250, 252
unisulcata, Campylaspis . . 250, 252, 263
Upupa . 37, 42
Uramba ..... 205, 239
uraniponnica, Finmarchinella . . 83, 86
valleculata, Campylaspis. 250, 254, 282-284
vallombrosae, Armadillidium . . .212
vanderhorsti, Haplochromis . . .162
vannus, Euplotes . . . . . 101
variabilis, Euplotes . . . . .124
variabilis, Porcellionides .... 204
velifer, Haplochromis . . . -143
venetus, Protracheoniscus . . . 207
Venezillo ...... 232
venustus, Armadillo . . . . .232
venustus, Venezillo .... 232
verecundus, Eptesicus . . . -134
verecundus, Philetar brachypterus 134-135, T 37
verrucosa, Campylaspis . . 250, 255, 297
verrucosus, Armadillo . . . .227
verrucosus, Diploexochus . . . .227
versicolor, Armadillidium . . . .212
Vespertilio . . . . 134, 172, 174
vespertilio, Porcellio .... 203
Vespertilionidae 73-82, 167-188, 317, 318, 319,
320, 322, 324, 325, 326, 327,
328, 329, 330
violaceus, Porcellio .... 198, 204
virgatus, Metoponorthus .... 197
viridis, Metoponorthus .... 197
viticola, Armadillo . . . . .221
vitrea, Campylaspis . . 250, 252, 263, 285
vittatus, Periscyphis . . . .217
vizzavonensis, Porcellio lugubris . . 201
vizzavonensis, Porcellio orarum . .201
vulcani, Tilapia .... 148, 149
vulgare, Armadillidium . . 210,212,221
vulgaris, Sturnus . . . . -37
wagleri, Psarocolius .
weberi, Armadillo
weberi, Spherillo
West, B. J. .
Whale-lice
White-beaked dolphin
37
. 231
231
95-102, i PI.
59-64, i PI. ; 65-72
59-64, i PI.
INDEX
wilsmorei, Armadillo
wilsmorei, Cubaris .
wingatii, Haplochromis
wintoni, Laephotis .
wintoni, Laephotis wintoni
wintoni angolensis, Laephotis
wintoni wintoni, Laephotis
woodi, Scotoecus
woodi, Scotoecus albofuscus
Xestadillo
. 221
. 221
162, 163
73. 74-75, 76, 77,
78, 79, 80, 81
73
73
73
176, 182, 183
175, 176-177
230
yemenensis, Porcellio
zachvatkini, Hemilepistus
zealandicus, Porcellio
zebricolor, Armadillo
zebricolor, Cubaris .
zenkewitchi, Euplotes
zigzag, Armadillo
zigzag, Diploexochus
zwartbergensis, Diploexochus
349
204
194
204
221
221
I2 4
227
227
227
17
ANATOMY OF HEAD AND NECK
IN THE HUIA (HETERALOCHA
ACUTIROSTRIS) WITH
COMPARATIVE NOTES ON OTHER
CALLAEIDAE
P. J. K. BURTON
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 27 No. i
LONDON: 1974
FRONTISPIECE. Male and female Huias (Heteralocha acutirostris) showing probable feeding
methods. The male is depicted excavating decayed wood by the 'gaping' technique described
in the text.
17 JULI9I
ANATOMY OF HEAD AND NECK IN THE
HUIA (HETERALOCHA ACUTIROSTRIS) WITH
COMPARATIVE NOTES ON OTHER CALLAEIDAE
BY
PHILIP JOHN KENNEDY BURTON
Pp 1-48 ; i Plate, 26 Text-figures
BULLETIN OF
THE BRITISH MUSEUM (NATURAL HISTORY)
ZOOLOGY Vol. 27 No. I
LONDON: 1974
THE BULLETIN OF THE BRITISH MUSEUM
(NATURAL HISTORY), instituted in 1949, is
issued in five series corresponding to the Departments
of the Museum, and an Historical series.
Parts will appear at irregular intervals as they
become ready. Volumes will contain about three or
four hundred pages, and will not necessarily be
completed within one calendar year.
In 1965 a separate supplementary series of longer
papers was instituted, numbered serially for each
Department.
This paper is Vol. 27, No. i, of the Zoological series.
The abbreviated titles of periodicals cited follow those
of the World List of Scientific Periodicals.
World List abbreviation :
Bull. Br. Mus. nat. Hist. (Zool.)
Trustees of the British Museum (Natural History), 1974
TRUSTEES OF
THE BRITISH MUSEUM (NATURAL HISTORY)
Issued 13 June, 1974 Price 2-30
ANATOMY OF HEAD AND NECK IN THE
HUIA (HETERALOCHA ACUTIROSTRIS) WITH
COMPARATIVE NOTES ON OTHER CALLAEIDAE
By PHILIP J. K. BURTON
CONTENTS
Page
I. INTRODUCTION ......... 3
II. METHODS AND MATERIAL ....... 5
III. SKULL AND LIGAMENTS ........ 6
IV. JAW MUSCULATURE ........ l6
V. BUCCAL CAVITY AND GLANDS ....... 24
VI. TONGUE APPARATUS ........ 24
VII. NECK AND NECK MUSCULATURE ...... 28
VIII. OTHER CALLAEIDAE ........ 34
IX. FUNCTIONAL ASPECTS ........ 36
X. SEXUAL DIMORPHISM ........ 40
XI. CONCLUDING REMARKS ........ 44
XII. ACKNOWLEDGEMENTS ........ 46
XIII. REFERENCES ......... 46
SYNOPSIS
Spirit specimens and skeletons of the Huia (Heteralocha acutirostris) have been used in a
detailed study of feeding adaptations in this extinct species. The Huia showed remarkable
sexual dimorphism in bill form, and particular attention is paid throughout to differences
between male and female. Comparison is made with the other members of the Callaeidae -
Creadion carunculatus and Callaeas cinerea. The features studied indicate that the Huia was
highly specialized for feeding by 'gaping' in decayed timber, in order to gain access to beetle
grubs. The full extent of sexual dimorphism revealed by this study is reviewed, and its sig-
nificance is discussed.
I. INTRODUCTION
THE extinct Huia (Heteralocha acutirostris) of New Zealand is frequently quoted as
an example of extreme sexual dimorphism among birds. This dimorphism is shown
most conspicuously by the bill, which in the male was typically fairly straight,
moderately long, and tapered evenly to a point ; while in the female it was much
longer, more slender, and strongly downcurved. There is little information on the
extent of dimorphism beyond this. Phillipps (1963) mentions plumage differences
and a tendency towards stronger claws in older males, and Selander (1966) gives
bill, wing and tarsus measurements for a small sample of each sex. The anatomical
study of the Huia by Garrod (1872) concentrated principally on establishing the
affinities of the species, and did not inquire closely into its sexual dimorphism. The
problem of its relationships is still not fully resolved beyond the fact that the Huia
is closely related to two other New Zealand species -the Saddleback (Creadion
4 P. J. K. BURTON
camnculatus) and the Kokako (Callaeas cinerea) (Stonor, 1942). The three together
form the family Callaeidae, placed between the Dicruridae and Grallinidae by Mayr
and Green way (1956) ; close affinity with the Sturnidae has been suggested in the
past (Garrod, 1872 ; Gray, 1870).
The present investigation has been concerned with the anatomical basis of the
Huia's feeding specializations. In particular, I have attempted to discover the
full extent of sexual dimorphism in feeding structures, and to correlate this where
possible with bill form and feeding methods. It is thus a study of adaptation,
unusual in that the principal forms to be compared are the two sexes of a single
species. The material on which the investigation is based is contained in the avian
anatomical collections of the British Museum (Natural History). These consist of
six spirit and one osteological specimens of the Huia, one spirit specimen of the
Saddleback and four spirit specimens of the Kokako. It is thought that the Huia
spirit specimens may be the only fluid-preserved examples of this species in existence,
although there is at least one skeleton in New Zealand (Phillipps, 1963) and another
in the U.S.A. (Bock, igGoa.).
The history of the Huia's disappearance, and many aspects of its life and habits,
are summarized by Phillipps. Last reported alive in 1907, it is probable that some
survived after this date, and Phillipps entertains the possibility that a few may still
exist. Its range was restricted to the mountainous Wellington Province of North
Island, New Zealand. Here, the combination of a mild climate and heavy rainfall
has produced densely forested country with a high incidence of timber decay and
epiphytic growth. The Huia fed principally on insects, varied with some vegetable
matter, but by far the most important prey appears to have been the larvae of the
Huhu beetle, Prionoplus reticularis (Cerambycidae) . This large beetle is common
in many parts of New Zealand, and its larvae, found in decaying timber, may reach
a length of 70 mm and a width of 20 mm. The most detailed account of the Huia's
manner of extracting these is provided by Buller (1888). This account has been
often quoted, but because of its relevance to the present study, it seems essential
to do so again. Buller's description refers to a pair of captive Huias which at first
he fed on individual huhu grubs :
'On offering one of these to the Huia he would seize it in the middle, and, at once transferring
it to his perch and placing one foot firmly upon it, he would tear off the hard parts, and then
throwing the grub upwards to secure it lengthwise in his bill, would swallow it whole . . . They
seemed never to tire of probing and chiselling with their beaks. Having discovered that the
canvas lining of the room was pervious, they were incessantly piercing it, and tearing off large
strips of paper, till, in the course of a few days, the walls were completely defaced.
But what interested me most of all was the manner in which the birds assisted each other in
their search for food, because it appeared to explain the use, in the economy of nature, of the
differently formed bills in the two sexes. To divert the birds, I introduced a log of decayed wood
infested with the huhu grub. They at once attacked it, carefully probing the softer parts with
their bills, and then vigorously assailing them, scooping out the decayed wood till the larva or
pupa was visible, when it was carefully drawn from its cell, treated in the way described above,
and then swallowed. The very different development of the mandibles in the two sexes enabled
them to perform separate offices. The male always attacked the more decayed portions of the
wood, chiselling out his prey after the manner of some Woodpeckers, while the female probed
with her long pliant bill the other cells, where hardness of the surrounding parts resisted the
ANATOMY OF HEAD AND NECK IN THE HUIA 5
chisel of her mate. Sometimes I observed the male remove the decayed portion without being
able to reach the grub, when the female would at once come to his aid, and accomplish with her
long slender bill what he had failed to do. I noticed, however, that the female always appro-
priated to her own use the morsels thus obtained.'
An additional reference to the different feeding techniques of male and female is
the note by J. M. Wright quoted by Oliver (1955):
'I have watched them in pairs hunting for wetas.* The male would tear away at the outer
part of a green sapling. The female then tried to retrieve the weta with her long slender bill.
If not successful she would stand back while the male tried to enlarge the hole. Unless the
tree was a maire the birds would generally succeed and would then fly away with the weta. 1
Phillipps (1963) also provides the most detailed information available on the
extent of dimorphism ascertainable from skins, and on the range of variation in
the two sexes, based on 119 specimens in New Zealand museums. Young birds
show little difference in size and shape of the bill. Some male bills were more curved
than usual, but were normally deeper than those of females. The male bills reached
an extreme length of 59-60 mm (from feathers), and females up to 104 mm. Phillipps
mentions two doubtful cases of birds with bills 63 mm long which he has treated as
males, though they may be females. The skin collection of the British Museum
(Natural History) includes two doubtful individuals ; one, with a bill length of
76-4 mm is labelled as a male ; the other, labelled as a female, has a bill of 71-8 mm.
Excluding these, bill length data for other specimens in the collection may be
summarized as follows :
Males : Mean 59-7 ; Min. 54-7 ; Max. 65-0 (16 specimens).
Females : Mean 96-3 ; Min. 87-1 ; Max. 108-0 (7 specimens).
Buller (1878) also gives details on the range of bill form in the Huia and figures
several specimens, including a female with the upper jaw much longer than the lower,
recalling some species of Hemignathus (Drepanididae) .
II. METHODS AND MATERIAL
Most of the observations reported here were obtained simply by dissection using
a stereomicroscope at powers mainly in the range X2| to X20. The main com-
plicating factor was the irreplaceable nature of the Huia specimens, and because
of this, it was thought prudent to keep dissections of this species to a minimum. In
general, dissections have been limited to the left side, and as little structure as
possible has been removed. Because of its history, the single specimen of Creadion
carunculatus was regarded as of similar value, and no dissections were attempted
which would have required removing or severing structures.
For convenient reference, the Huia specimens were given a simple individual
coding in addition to their British Museum (Natural History) Registration numbers.
* Orthoptera, Gryllacridoidea.
6 P. J. K. BURTON
Details of the specimens of Callaeidae used, with their conditions at commencement
of the study, are as follows :
Heteralocha acutirostris
$ i. Reg. No. A 1973.1.3. Intact spirit specimen.
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ANATOMY OF HEAD AND NECK IN THE HUIA 15
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16 P. J. K. BURTON
sesamoids, one placed laterally and the other posterior to the quadrate. In common
with all passerines so far studied, the lower jaw has no medial brace (see Bock,
I96ob).
IV. JAW MUSCULATURE
A work of major importance for studies of avian jaw musculature is Lakjer's
(1926) review of the trigeminal musculature. For passerines in particular, the
paper by Fiedler (1951) provides valuable clarification. Investigations by Bock
(i96oa and MS) have resolved many remaining problems of structure and ter-
minology within the order, and his nomenclature for the subdivision of complex
muscles is the one followed here. The jaw musculature of the Callaeidae remains
undescribed beyond the comments of Garrod (1872) and Lowe (1938) on the large
size of the 'digastric muscle' (= M. depressor mandibulae).
In describing the structure of complex muscles it is often useful to designate their
various aponeuroses by letters or numbers. It is far from clear to what extent these
aponeuroses can be homologized between different groups, although the work of
Starck and Barnikol (1954) has made progress in this respect for M. adductor
mandibulae externus. In the present case, a simple numbering system has been used
where necessary, but is not intended to imply homology with any numbered apo-
neuroses described from other species, with the exception of Aps. i, 2 and 3 in M.
adductor mandibulae externus.
M. adductor mandibulae externus
This important and complex muscle acts to raise the lower jaw and to maintain
a grip on objects held in the bill. Its architecture is intricate, and the relationships
of its aponeuroses can be better understood by the examination of cross sections
(Fig. 18). The abbreviation 'M.a.m.e.' is used in parts of the description to shorten
the otherwise unwieldy names of some subdivisions. The three major aponeuroses
described by Starck and Barnikol (1954) in a wide range of birds of several orders
can be recognized with little difficulty in Heteralocha, and their numbering here
(Aps. i, 2 and 3) is the same as Starck and Barnikol's.
M . adductor mandibulae externus rostralis
The most dorsal part of M. adductor mandibulae externus, with an extensive
fleshy origin on the cranium. Insertion is made principally via Ap. i, which nar-
rows anteriorly to form a flat tendon attached to a crista on the surangular. Follow-
ing Bock, three sections are recognized :
M.a.m.e. rostralis medialis. Origin is from the postero-lateral border of the orbit,
medial to the postorbital process, and from a vertically oriented aponeurosis (Ap. 4)
attached to the orbit along the medial border of the muscle. Insertion is made via
Ap. i. Medially, a group of fibres from the ventral part of the muscle overlaps part
of Ap. i as a more or less distinct slip.
M.a.m.e. rostralis temporalis. Origin is fleshy from the border of the temporal
fossa ; a short aponeurosis (Ap. 5) runs forwards from the tip of the postorbital
ANATOMY OF HEAD AND NECK IN THE HUIA
Mdt
Mdm
L po
FIG. ii. Heteralocha acutirostris, male (<$ i), jaw muscles and ligaments.
For abbreviations see p. 47.
process across the dorsal lateral surface, and in some specimens extends onto the
dorsal surface of M.a.m.e. rostralis medialis. The fibres of this section converge
anteriorly, falling into an ill-defined bipinnate arrangement, and are attached to the
ventro-lateral surface of Ap. i ; a few fibres overlap its dorsal surface posteriorly, but
it is exposed for some 8 mm from the insertion.
M.a.m.e. rostralis lateralis. This section lies superficial to the dorsal part of
M.a.m.e. ventralis. Its origin is from the lateral edge of the zygomatic process, and
from the lateral edge of Ap. 2 in its narrow posterior region. Fibres run upward
and medially from the origin and insert on the lateral surface of the mandible over
a narrow zone dorsal to the insertion of M.a.m.e. ventralis. This insertion is fleshy,
and via a superficial aponeurosis (Ap. 6) which merges medially with Ap. i. The
anterior ventral part of this section lies medial to the jugal bar and is in consequence
flattened into a relatively thin sheet of fibres.
M. adductor mandibulae externus ventralis
Origin is from the zygomatic process, by a strong aponeurosis (Ap. 2) which
traverses the gap between the cranium and mandible as a fairly narrow tendon, then
fans out over the lateral surface of the muscle. Fibres arising from this aponeurosis
make an extensive fleshy insertion on the lateral surface of the mandible, spanning
the fenestra in the posterior part of the mandible.
M. adductor mandibulae externus caudalis
The shortest section of M. adductor mandibulae externus. Origin is from the
otic process of the quadrate and from the ventral surface of the posterior half of
Ap. 2. The origin is fleshy, and by a weak aponeurosis (Ap. 7). Insertion is on the
dorsal lateral edge of the surangular, posterior and ventral to that of Ap. i. The
insertion is made by a short, narrow aponeurosis (Ap. 3), which bifurcates within
i8
P. J. K. BURTON
the body of the muscle to form a dorsal (Ap. 3a) and a ventral (Ap. 3b) branch.
Ap. 7 passes between 3a and 3b before fading out. The muscle shows multipinnate
structure, clearest in its middle third, and most conspicuous about Ap. 3a. Fibres
run forwards from Ap. 2 to Ap. 3a, back from Ap. 3a to Ap. 7, and forwards again
from Ap. 7 to Ap. 3b.
M. pseudotemporalis superficialis
A muscle of relatively simple structure, whose action is adduction of the lower jaw.
The origin is a broad fleshy one on the posterior wall of the orbit, immediately
medial to M.a.m.e. rostralis medialis. There is also a weak dorsal aponeurosis
attached to the orbit. Insertion is made via a strong tendon attached to the medial
side of the surangular, ventral to the ramus mandibularis of the trigeminal nerve.
Mdt
Md m
L po
FIG. 12. Heteralocha acutirostris, female ($ C), jaw muscles and ligaments.
For abbreviations see p. 47.
M. pseudotemporalis profundus
This muscle is an adductor of the lower jaw, and by its action on the quadrate it
also functions to lower the upper jaw. It takes its origin from the dorso-lateral
surface of the orbital process of the quadrate, and from a strong aponeurosis attached
to the expanded tip of the orbital process. It makes an extensive fleshy insertion
on the medial surface of the mandible, overlapping slightly onto its dorsal edge,
adjacent to Ap. i of M.a.m.e. rostralis.
M. adductor mandibulae posterior
The actions of this muscle are similar to those of M. pseudotemporalis profundus.
Due to its situation closer to the mandible-quadrate articulation, it has a lower
mechanical advantage, but its upward component may be of value in holding the
mandible against the quadrate. M. adductor posterior is contiguous with M.
ANATOMY OF HEAD AND NECK IN THE HUIA
Mptvm Mptdm Mpsp Mpq1 Mamerm Mamert
M a m ev
M amec
FIG. 13. Heteralocha acutirostris (<$ i), jaw muscles in lateral view.
For abbreviations see p. 47.
pseudotemporalis profundus at its origin, which is also a fleshy one on the quadrate,
occupying the base of the orbital process and the dorsal surface of the quadrate body.
The two muscles are separated (following the criterion used by Lakjer, 1926) by
N. pterygoideus. The muscle inserts on the expanded region of the dorsal surface
of the mandible just anterior to the base of the internal process, and posterior to the
insertion of M.a.m.e. caudalis. The insertion is fleshy and via a weak dorsal apo-
neurosis.
M. pterygoideus
This important muscle complex acts both to lower the upper jaw and to raise the
lower. Its attachments to the palatine and pterygoid are arbitrarily regarded as the
origin, and attachment to the lower jaw as the insertion.
M. pterygoideus dorsalis later alis
This section has an extensive fleshy origin on the dorsal surface of the palatine.
It inserts on the medial surface of the mandible, posterior to M. pseudotemporalis
profundus, fleshily, and by a superficial aponeurosis (Ap. i), which is strongest
ventrally.
M. pterygoideus ventralis lateralis
Origin is from an aponeurosis (Ap. 2) which is attached to the postero-lateral tip
of the palatine, and extends for some distance across the ventral surface of the muscle.
The medial edge of this aponeurosis serves to demarcate it from M. pterygoideus
20 P. J. K. BURTON
ventralis medialis seen in ventral view. M. pterygoideus ventralis lateralis is at-
tached at the base of the internal process of the mandible, ventral to the insertion
of M. pterygoideus dorsalis lateralis. There is no overlap (Venter externus') onto
the ventral edge or lateral surface of the mandible as in many birds, since the ventral
medial edge of the mandible in this region is occupied by a forward extension of M.
depressor mandibulae.
M pt v m
M pt d m a
M pt d m p
Mpt r
Mptdl
Aplame Mpsp Mamerm
Mamert
5mm
FIG. 14. Heteralocha acutirostris ( i), jaw muscles in dorsal view.
For abbreviations see p. 47.
M. pterygoideus dorsalis medialis
In dorsal view, this muscle is seen to lie immediately adjacent and posterior to
M. pterygoideus dorsalis lateralis, separated from it by a narrow groove which widens
medially, exposing a small area of the dorsal surface of M. pterygoideus ventralis
medialis. Its origin is confined to the pterygoid, and occupies much of its surface.
The muscle falls into anterior and posterior portions, whose fibres diverge in a
posterior direction roughly along the lines of the pterygoid, so that it appears bi-
pinnate in dorsal view.
M. pterygoideus dorsalis medialis anterior originates more laterally on the ptery-
goid, and inserts narrowly on the mandible, immediately caudal to M. pterygoideus
dorsalis lateralis. The insertion is fleshy and aponeurotic, the aponeurosis (Ap. 3)
being strongest ventrally.
M. pterygoideus dorsalis medialis posterior originates medially on the pterygoid,
and fans out to a wide fleshy insertion on the dorsal part of the internal process of
the mandible, dorsal to that of M. pterygoideus ventralis medialis.
ANATOMY OF HEAD AND NECK IN THE HUIA
M pt v m
Eus
Mpt r
Ap 3a d m
M d m
5mm
FIG. 15. Heteralocha acutirostris (<$ i). M. pterygoideus and M. depressor in ventral view.
Left side of palate left intact to show papillae. For abbreviations see p. 47.
M . pterygoideus ventralis medialis
There is an extensive fleshy origin on the ventral surface of the palatine and
insertion is made on the internal process of the mandible, fleshily, and by a strong
dorsal aponeurosis (Ap. 4). The fibres of this section are orientated more nearly
parallel to the skull axis than those of any other part of M. pterygoideus. On the
medial side of the muscle is a slip (M. pterygoideus retractor) exclusively specialized
for retracting the palatal apparatus, attached at one end to the medial caudal tip
of the palatine, and at the other to the basitemporal plate of the skull. A few of its
most medial fibres are attached to the edges of the opening of the Eustachian tube.
M. protractor quadrati et pterygoidei
This muscle acts to raise the upper jaw by moving the palatal framework and
jugals forwards. It has a wide fleshy origin from the posterior part of the interorbital
septum and the adjacent region of the posterior orbital wall. The lateral border
of the muscle lies immediately adjacent to M. pseudotemporalis superficialis.
22 P. J- K. BURTON
Two parts can be distinguished. The more anterior and medial part (M. protractor
i), arising mainly on the interorbital septum, shows a bipinnate fibre arrangement.
Its raphe is a strong aponeurosis attached to a spur on the posterior dorsal tip of the
pterygoid, immediately adjacent to the quadrate. This raphe serves as the principal
site of insertion for the fibres of M. protractor i.
The lateral part (M. protractor 2) originating from the posterior wall of the orbit
inserts fleshily, and by a weak dorsal aponeurosis on the caudal edge of the basal half
of the orbital process.
Mptdl MptdmaMpss Mpql Mamerm
M pt d m p
/
Ma mert
M d m
Mamev Ap2ame Mamp Mamec
5mm
FIG. 16. Callaeas cinerea, jaw muscles in lateral view. For abbreviations see p. 47.
M. depressor mandibulae
This muscle, which depresses the lower jaw, is of enormous bulk (and, presumably,
fibre number) in Heteralocha relative to the other jaw muscles. There is a wide
fleshy origin on the squamosal, parietal and exoccipital. Fibres originating high on
the cranium converge to insert fleshily near the extremity of the long retroarticular
process, mainly on its medial side. Those originating lower, on the exoccipital, fan
out to insert fleshily on both surfaces of the retroarticular process. Fibres inserting
medially extend far forward beyond the retroarticular process along the ventral half
of the mandible, ventral to the insertion of M. pseudotemporalis profundus. This
extension is here termed the pars anterior of M. depressor mandibulae.
ANATOMY OF HEAD AND NECK IN THE HUIA
Mptdl Mptdma Mptdmp Mpq1 Mpss
M a merm
M a m e v
M a m e rt
Mdm
5mm
FIG. 17. Callaeas cinerea, jaw muscles in dorsal view. For abbreviations see p. 47. M.
pseudotemporalis profundus is concealed by M. pseudotemporalis superficialis.
2mm
FIG. 18. M. adductor mandibulae externus, TS at about the midpoint of the otic process of
the quadrate, and at right angles to it. Numbering of aponeuroses as explained in text.
A - Callaeas cinerea. B - Heteralocha acutirostris (<$ i). The left side is lateral in both
diagrams.
The following aponeuroses can be distinguished :
Ap. i. This is attached to the cranium around the dorsal edge of the origin on the
squamosal and parietal, and extends some way across the lateral surface
of the muscle.
Ap. 2. A strong aponeurosis with a narrow attachment at about the middle of the
posterior border of the auditory meatus, fanning out across the lateral
surface of the muscle, and acting as a surface of origin for fibres inserting
on the anterior half of the lateral surface of the retroarticular process.
24 P. J- K. BURTON
Ap. 3. A strong aponeurosis attached to the occipital crest, which marks the
postero- ventral limit of the origin. Ap. 3 extends some way across the
medial surface of the muscle, and becomes stronger on the exoccipital
process, whence it is produced as a strong raphe (Ap. 3a) across the
ventral region of the muscle, and on into the pars anterior. Fibres
arising from its medial side insert mainly on the medial surface of the
internal process of the mandible. Those running from its lateral side
insert on the medial surface of the basal half of the long retroarticular
process, on the posterior face of the internal process at its base, and on
the ventral medial surface of the mandible.
Ap. 4. An aponeurosis attached to the dorsal edge of the internal process of the
mandible, extending across much of the dorsal surface of the short part
of the muscle between this and the exoccipital. This aponeurosis is
branched and infolded within the muscle.
Ap. 5. An aponeurosis attached to the dorsal edge of the retroarticular process,
but concealed from lateral view by the thin sheet of fibres originating
from Ap. 2, and inserted on the lateral surface of the process.
V. BUCCAL CAVITY AND GLANDS
Backwardly directed horny papillae are abundant on the surface of the palate.
They are longest where they fringe the choanae, and in a densely clustered transverse
row posterior to the internal opening of the Eustachian tubes. The surface of the
larynx is also papillate, the papillae being longest at its posterior margin.
Three pairs of salivary glands are present. The Gl. angularis oris lies on the side
of the head immediately below the skin and just ventral to the jugal bar ; its duct
opens at the angle of the gape. A second gland, which may be termed a Gl. palatinae
(see Antony, 1920), lies between M. depressor mandibulae, pars anterior, and M.
pterygoideus ventralis lateralis, immediately below the mucosa of the palate. The
third gland is a sublingual one. It lies immediately lateral to M. genioglossus, and
extends from the level of the anterior border of M. mylohyoideus to the region of
the tongue base. Its duct runs alongside M. genioglossus, but diverges from it
near the mandibular symphysis to open in a lateral position.
VI. TONGUE APPARATUS
The tongue is narrow, tapering evenly towards its anterior tip which is somewhat
frayed and brush like (Fig. 19). The lateral edge bears a few papillae posteriorly,
and the posterior edge is crowded with pointed papillae, longest laterally. Tongue
length for the specimens examined is shown in Table i.
The tongue skeleton is similar to that of many passerines. The paraglossalia
which provide support for the tongue itself have long posterior processes, and
anteriorly they meet and run side by side to a point about one-third of the way from
the tip of the tongue. The remainder of the tongue is purely corneous. The basihyal
ANATOMY OF HEAD AND NECK IN THE HUIA
5mm
FIG. 19. Heteralocha acutirostris (<$ i), tongue and larynx in dorsal view.
is a slender rod of roughly triangular section, with the apex of the triangle upper-
most. The flattened urohyal, expanded at its posterior tip, and the hyoid horns
(each consisting of the ceratobranchial posteriorly and epibranchial distally) are of
similar form to those of many passerines.
The tongue muscles in Heteralocha show few unusual features, and no significant
sexual dimorphism was encountered in the dissections. The descriptions given are
therefore brief ones ; illustrations are given in Figs. 21 and 22. More detailed
accounts and background information are given in the general review by George and
Berger (1966) and the paper on passerine tongue muscles by Engels (1938). A very
full description of the tongue muscles of a single passerine species is that by Bock
(1972) for the extinct Ciridops anna (Drepanididae) . For discussions on function,
reference may also be made to the account of wader tongue muscles by Burton
(I974)-
M. mylohyoideus
A thin muscular sheet with a long narrow origin on the medial side of the man-
dibular ramus, inserting on a median raphe. M. mylohyoideus lies ventral to the
tongue and all the muscles attached to it.
M . serpihyoideus
Origin is on the occipital plate, medial to the exoccipital process (Bock, igGob,
p. 38) and insertion is on a median raphe continuous with that of M. mylohyoideus.
26
P. J. K. BURTON
M . genioglossus
A narrow, strap-like muscle whose origin is from the posterior edge of the mandi-
bular symphysis. The left and right muscles lie side by side near the origin and then
diverge, running along the ventral side of the mucosa of the floor of the buccal
cavity, on either side of the position of the tongue. In the region of the basihyal,
the fibres of the muscle fan out to insert on the connective tissue and mucosa over-
lying the basihyal and its musculature, and, anteriorly, on the posterior process of
the paraglossa.
FIG. 20. Callaeas cinerea, tongue and larynx in dorsal view.
M. stylohyoideus
A long, narrow muscle originating on the ventral edge of the base of the retro-
articular process, and running along the anterior edge of M. serpihyoideus, to insert
on the lateral surface of the basihyal, just anterior to M. thyreohyoideus.
M. branchiomandibularis (= M. geniohyoideus of many authors)
A bulky muscle whose origin is on the medial surface of the mandible, ventral
to that of M. mylohyoideus. A broad anterior and narrow posterior position can be
distinguished. The two run parallel to insert on the hyoid horn. The anterior
portion meets the horn from the ventral side, and is twisted around it for some dis-
tance before inserting on the epibranchial. The posterior portion meets the anterior
on the dorso-medial side, and merges with it.
M. ceratohyoideus
A thin, weakly developed muscle which was found only in <$ i. It originates on
the hyoid horn, on the ventral lateral surface of the distal tip of the ceratobranchial,
ANATOMY OF HEAD AND NECK IN THE HUIA
Apm
Mthh
Mseh
FIG. 21. Heteralocha acutirostris ($ B), tongue muscles in ventral view.
For abbreviations see p. 47.
deep to M. ceratoglossus. The insertion is on a median raphe continuous with that
of M. serpihyoideus and M. mylohyoideus, near the posterior end of the urohyal.
M. ceratoglossus
A unipinnate muscle whose fibres arise on the surface of the ceratobranchial, and
the anterior end of the epibranchial, and insert on a long lateral tendon which is
itself inserted on the ventral surface of the paraglossal, just level with the anterior
tip of the basihyal. No fibres insert on the tendon over the region lying alongside
the basihyal, but just anterior to its attachment to the paraglossal, a small fleshy
slip arises. It merges with that from the other side, and together they insert on a
strong medial aponeurosis which runs along the ventral side of the paraglossalia, and
inserts on the corneous anterior part of the tongue. This slip is referred to by Bock
(1972 and MS) as M. hypoglossus anterior. A similar slip in shorebirds was described
by Burton (1974) as M. ceratoglossus anterior ; the term 'M. hypoglossus anterior' is
used by Burton (1974) to refer to a quite distinct muscle, apparently absent from
28 P. J. K. BURTON
passerines, arising on the posterior tip of the paraglossals and also inserting on the
median aponeurosis.
M. hypoglossus obliquus
Origin is on the postero-lateral process of the paraglossa. The right and left
muscles are merged, the whole forming a bulky loop passing ventral to the anterior
third of the basihyal.
Mthh Mtrh
M th h M tr I
FIG. 22. Heteralocha acutirostris (? C), ventro-lateral view of tongue muscles attached to the
cricoid cartilage. Cricoid and urohyal shown stippled. Both left and right Mm.
tracheolateralis are visible. For abbreviations see p. 47.
M. tracheohyoideus
Origin is from the skin of the neck, and insertion on the lateral surface of the
cricoid, just below the dorsal origin of M. thyreohyoideus.
M. tracheolateralis
The muscle originates on the syrinx, and passes along the side of the trachea,
broadening anteriorly to insert by two heads on the lateral surface of the cricoid.
M. thyreohyoideus
This muscle originates from the lateral surface of the cricoid, by a dorsal and a
ventral slip. The dorsal one is slender, and is attached just below the dorsal edge of
the cricoid. The ventral slip is broader, and arises between the two heads of insertion
of M. tracheolateralis. These slips unite anterior to the larynx and insert on the
anterior lateral surface of the basihyal posterior to the insertion of the more slender
M. stylohyoideus.
M. ceratoglossus superior (Bock, 1972) was not found in Heteralocha. The glottal
muscles (M. thyreoarytenoideus and M. constrictor glottidis) conform to Bock's
description for Ciridops anna.
VII. NECK AND NECK MUSCULATURE
The thorough review by Boas (1929) has provided the basis for most subsequent
studies of the avian neck and its musculature. Boas concentrated on non-passerines,
principally large species, but detailed information on several small passerines is
given by Palmgren (1949). There is, however, little information on the cervical
ANATOMY OF HEAD AND NECK IN THE HU1A
Mrcl M co M s co M sp
Mil
Mrcv Mfcb Mas 6-3 Mas 7-3 Mas 8-5
10mm
FIG. 23. Heteralocha acutirostris ($ C), lateral view of superficial neck muscles.
For abbreviations see p. 47.
muscles of larger passerines comparable in size with the Callaeidae. The majority
of neck muscles perform several actions which interact in a complex way, and their
individual functions are omitted from the present account in the interest of brevity.
Reference may be made to Boas and Palmgren, and to the excellent discussions on
neck muscle function by Zusi (1962, 1969).
There are 14 cervical vertebrae in Heteralocha, as in most passerines. This total
includes the two cervico-dorsal vertebrae (13 and 14) which bear movable ribs not
articulating with the sternum, but from a functional standpoint are best treated
with the neck. The rib on 13 is extremely short, while that on 14 is long, almost
reaching the sternum, but lacking an uncinate process. Six pairs of ribs actually
articulate with the sternum, as in Cor cor ax, but unlike most passerines which only
have 5 ribs attached to the sternum (Beddard, 1898).
Boas showed that the neck of birds consists of three sections, distinguishable by
both functional and morphological differences. Section I, the most anterior, can
only be flexed downward, and Section II only upward. Section III can be flexed
downward, and also upward at its anterior end. In Heteralocha, the constitution of
the segments agrees with Palmgren's division for smaller passerines (although Palm-
gren omitted the cervico-dorsal vertebrae from his count). The division is as
follows : Section I, vertebrae i to 4 ; Section II, vertebrae 5 to 9 ; Section III,
vertebrae 10 to 14. The vertebrae of Section I (except the atlas) have strong neural
spines and hypapophyses. Those in Section II lack neural spines (except for a weak
one on 5) and have no hypapophyses ; they are more elongated than the vertebrae
of Sections I and III. The vertebrae of Section III have hypapophyses ; that on 10
is weak, but they increase in size posteriorly. The last, and largest, hypapophysis
is on 15, the first dorsal vertebra. Vertebra 14 has a strong neural spine.
The account below follows the same order and terminology as Palmgren, with
modification in the case of Mm. splenii accessorii and Mm. intercristales.
30 P. J. K. BURTON
M. biv enter
This muscle arises from the dorsal surface of the aponeurosis of origin of M. spinalis
in the region of 13, and inserts on the dorsomedial edge of the occipital deep to M.
complexus. The muscle consists of two fleshy bellies linked by a flat tendon ex-
tending approximately from 8 to 5. Little variation was encountered.
M. spinalis
The muscle consists of a series of fleshy slips from the ventral surface of an
aponeurosis attached to the neural spines of vertebrae 14 to 18. These insert on the
anapophyses of 2 and of 5 or 6 to 13 ; the most posterior of these is feeble and in-
distinct in most of the specimens. Variations in the specimens of Heteralocha were
as follows :
Slips to both 5 and 6 present : $ A, $ B.
Slip to 6 present, slip to 5 absent : ^ i, ^3, $ C.
Slip to 5 present, slip to 6 absent : $2.
Mm. splenii colli
These muscles are a series of slips arising from the lateral surfaces of the neural
spines of 3 or 4 successive vertebrae, and joining the most anterior slip of M. spinalis
which inserts on 2. The vertebrae of origin in the Heteralocha specimens are :
4 to 7 : <$ i, $'s A, B and C.
4 to 6 : $ 2, ^ 3.
Mm. splenii accessorii and Mm. inter cristales
In order to clarify the relationships between these two rather similar groups of
muscles it seems desirable to deal with them together. The muscles included under
these headings in Heteralocha are as follows :
a. A muscle running from the anterior surface of the neural spine of 14 to the
transverse-oblique crest of 13, and a similar muscle connecting 13 and 12.
b. A series of muscles connecting the transverse-oblique crests of successive verte-
brae from 13-12 to 6-5.
c. Muscles arising on the ventro-lateral surfaces of the neural spines of 5 to 3 and
inserting on the transverse-oblique crests of 4 to 2.
d. A muscle arising on the dorso-lateral surface of the neural spine of 3 and insert-
ing on the anapophysis of 2, and a similar but weaker muscle arising on the neural
spine and medial part of the neural arch of 2 and inserting on the anapophysis of i.
That arising on 3 closely resembles Mm. splenii colli.
e. A narrow, flat slip arising from the ventral part of the neural spine of 5 and
inserting on the anapophysis of 3, somewhat resembling Mm. dorsales pygmaei.
According to Palmgren's criteria, groups a and b only should be regarded as com-
prising the Mm. intercristales, and groups c, d and e should be treated as Mm. splenii
accessorii. (In the small species examined by Palmgren the two muscles running
antero-laterally from the neural spine of 3 are inseparable.) However, it would
seem more consistent with Boas's work to include group c also in Mm. intercristales,
and to reserve the term Mm. splenii accessorii for groups d and e.
ANATOMY OF HEAD AND NECK IN THE HUIA 31
No noteworthy variations between individuals were found.
M. splenius capitis
This muscle originates from the neural spine of 2 and inserts on the posterior
surface of the skull deep to M. complexus and M. bi venter cervicis. No variations in
siting were found, and there is little indication of cruciform structure (Burton,
i97ia).
1
2
3
4
5
6
7
To 15
FIG. 24. Diagram to show arrangement of slips of some ventral muscles in the anterior
part of the neck in Heteralocha acutirostris (? B). Heavy lines = M. flexor colli brevis.
Broken lines = M. flexor colli profundus. Fine lines = anterior slips of M. longus colli
ventralis.
Mm. Pygmaei
Origin is from the medial region of the neural arches of 12 to 8. Each muscle
inserts on the lateral edges of the transverse-oblique crest of the second vertebra
anterior to it except that from 12 which inserts on n. The latter muscle is weak in
most of the specimens and absent in $ i.
Mm. inter spinales
These three muscles connect the well-developed neural spines of 2, 3, 4 and 5, and
were present in all specimens examined.
Mm. ascendentes cervicis
These muscles arise on the diapophyses of cervical vertebrae up to and including 6.
Most consist of two slips inserting on the anapophyses of the second and third
vertebrae anterior. That from 7 consists of three slips inserting on 3, 4 and 5 in all
specimens, and that from 8 sends an additional slip to 4 in ^'s i and 2. The short
slips arising on 12 and posteriorly are relatively weak, and also showed some minor
variations in siting. The series is continued posteriorly as Mm . ascendentes thoracicis,
but those arising posterior to 14 were not dissected.
Mm. intertransversarii
These laterally situated muscles connect successive vertebrae. The most anterior
are those from 3 to 2, and the most posterior those from 13 to 12. Each muscle
arises from the anterior surface of the transverse process and inserts on the posterior
surface of the transverse process of the vertebra in front ; that from 4 to 3 inserts on
32 P. J. K. BURTON
the medial surface of the rib of 3, and that from 3 to 2 inserts on the lateral surface
of the centrum of 2.
As far as 6-5, the Mm. inclusi lie deep to Mm. intertransversarii, and closely
associated with them. Palmgren regards the muscles anterior to this as continuing
the series of Mm. inclusi ; Boas and Zusi treat them with Mm. intertransversarii as
is done here.
The Mm. intertransversarii are multipinnate muscles, traversed by interdigitating
raphes from origin and insertion. The muscles are bulkiest and the number of
raphes greatest in the region from 7 to 10. Up to 9 raphes have been detected in
9-8 or in 10-9, the number and situation of raphes showing small individual varia-
tions. The muscle connecting 13 and 12 is reduced to a small dorsal slip, while
those anterior to 5 are also of small size.
Mm. inclusi
These muscles are concealed by Mm. intertransversarii, and can only be separated
from them with difficulty. Each one arises on the anterior surface of the transverse
process medial to M. intertransversarius, and inserts on the lateral surfaces of the
neural arch and centrum of the next vertebra in front. Most show division into
dorsal and ventral bellies (Mm. inclusi superiores and inferiores). In the most
posterior two (12 to n and n to 10), only inferiores can be distinguished. The
most anterior are those connecting 5 and 6.
M. longus colli ventralis
This complex muscle consists of a series of fleshy slips arising on the sublateral
processes, hypapophyses and anterior part of the centra of vertebrae 15 to 6. The
main part of the muscle inserts by a series of 7 tendons on the ribs of n to 5. Each
vertebra sends a slip to join each of the tendons traversing it ; there are thus 7 such
slips from each of vertebrae 15 to 12 after which the number of slips decreases by
one for each vertebra anterior to this. In the region 15 to 12 the slips are densely
crowded and difficult to separate and some of the deepest fibres appear to attach to
adjacent vertebrae.
A smaller group of slips situated anteriorly is also included with M. longus colli.
Three of these arise from a tendon attached to the sublateral process of 7, which
also provides origin for part of M. flexor colli profundus. They insert on ribs 3 and
4 by short aponeuroses and on the long tendon attached to rib 5. Slips also arise
on the ventral anterior surfaces of the lateral processes of 7, 6 and 5, the two former
in close association with the Mm. intertransversarii, and immediately below them.
These attach on rib 3, and those from 6 and 7 also on rib 4 ; minor variations occur.
Two slips arise from the sublateral process of 6 and insert on ribs 3 and 4, and there
is also a longer slip arising on 9 (8 in $ C) which inserts by a weak tendon on rib 4.
This tendon also appears to receive some fibres from the posterior part of the muscle.
M. flexor colli brevis
Lateral and medial parts may be distinguished in this muscle. The lateral part
constitutes the greater bulk of the muscle and originates from the ventral surface
ANATOMY OF HEAD AND NECK IN THE HUIA
33
of the lateral strut of 3 ; and from the lateral processes of 4 and 5 in the $'s and 6 also
in the g's. The medial portion is separated from the lateral by the anterior part of
M. longus colli ; it takes origin from the sublateral processes of 3, 4 and 5.
The lateral and medial portions join anterior to rib 3 and insert by a tendon on
the postero-ventral processes of the centrum of the atlas.
10 12 13 14 15
FIG. 25. Diagram to show arrangements of slips of some neck muscles in Heteralocha acu-
tirostris ($ A or $ B). Heavy lines = M. splenius capitis and Mm. splenii colli. Fine
lines = M. spinalis (dorsal) and M. longus colli ventralis (ventral). Broken lines = Mm.
splenii accessorii (as denned in text). Dotted lines = Mm. pygmaei. Tendons of M.
spinalis and M. longus colli ventralis are represented by double lines.
M. flexor colli profundus
Origin is from the sublateral processes of 4, 5, 6 and 7. The slips from 4 and 5
arise immediately deep to the medial portion of M. flexor colli brevis and insert on
the hypapophysis of 2. From 6, slips run to insert on the hypapophyses of 2, 3 and
4 (c? 2 > $ 3 $ B) or of 3 and 4 only ($ i, $ A, $ C). From 7 a long slip runs to the
hypapophysis of 2 ; this slip shares a tendinous origin with the medial slip of M.
longus colli from 7 to 3. In $ 2, $ 3 and $ B the slip bifurcates and a branch con-
tiunes forwards to fuse with M. flexor colli brevis near its insertion. The slips arising
on 6 show some fusion with M. rectus capitis ventralis at the origin.
M. complexus
Origin is from the lateral strut of 4, the diapophysis of 5, and from an aponeurosis
attached to the diapophysis of 6. Insertion is on the dorsal edge of the occipitals.
M. rectus capitis lateralis
Origin is from the hypapophyses of 2, 3 and 4, and insertion on the lateral dorsal
edge of the exoccipital.
M. rectus capitis superior
This muscle lies immediately superficial to M. flexor colli brevis. Origin is from
the lateral surface of neural arch i, from the anterior surfaces of anapophyses 2 and
34 P. J- K. BURTON
3, from the lateral strut of 4, and from the transverse processes of 5 and 6. Origin
from 6 is absent in all the female specimens and in < 2 ; origin from 5 is also absent
in $ C. Insertion is aponeurotic and fleshy on a ridge at the posterior edge of the
basitemporal plate.
M. rectus capitis ventralis
Origin is from the ventral surface of i, from the hyapophyses of 2, 3, 4 and 5, and
from the sublateral process of 6 where there is some fusion with M. flexor colli
profundus. The right and left Mm. recti capiti ventrales are fused in the midline
and insert together on the basitemporal plate, anterior to M. rectus capitis superior.
VIII. OTHER CALLAEIDAE
Creadion carunculatus
The description of the Huia's skull by Oliver (in Phillipps, 1963) refers to the
general similarity of the skull of Creadion. Oliver notes the greater extent of ossi-
fication in Heteralocha, manifested especially at the front of the orbit. The skull
and skeleton of this species are figured by Shufeldt (1913), who includes some brief
notes on its skull for comparison with Anthochaera carunculata (Meliphagidae) .
The following additional points may be noted. The retroarticular process is
highly developed in Creadion, but is nevertheless relatively shorter than that of the
Huia ; there is no raised occipital crest, and little development of an exoccipital
process. Otherwise, skull proportions are similar to Heteralocha, particularly
female specimens, though the quadrate and pterygoid are relatively smaller. The
skull is apparently larger relative to body size in the Saddleback if a rough index
(sternum length) from a single specimen can be relied upon. Ligaments have been
removed from the skull of the available specimen, and could not be studied.
With the exception of M. depressor mandibulae, jaw muscles have also been
removed from the specimen of Creadion. This remaining muscle is, however, of
considerable interest, since it possesses a pars anterior as in Heteralocha - a feature
which otherwise appears to be unique among birds so far studied. The rest of the
muscle is also well developed, and similar in structure to that of the Huia, though
relatively less massive. The tongue, like the jaws, is smaller relative to the skull in
Creadion, but the hyoid musculature is closely similar in both Saddleback and Huia.
Due to the historic interest of the specimen, only limited dissection of the neck
muscles could be undertaken for Creadion. Five of the muscles exhibiting variation
in the Huia or among other passerines were examined. These showed an overall
reduction in the number of sites of attachment. M. spinalis lacks the slip inserting
on 5 which is present in three of the Huia specimens, though a slip to 6 (absent in
Huia <$ 2) is present. M. splenius colli lacks the slip to 7 found in four of the Huia
specimens. M. flexor colli brevis and M. rectus capitis superior lack attachment to 6,
a feature found in three and two of the male Huia specimens respectively. M. corn-
plexus also lacks a slip to 6, although this is present in all six Huias dissected.
ANATOMY OF HEAD AND NECK IN THE HUIA 35
Callaeas cinerea
The skull and other features of the skeleton in the Kokako have been described in
some detail by Stonor (1942). It is sufficient here to draw attention to the extensive
differences in skull proportions and geometry between this bird, on the one hand,
and Heteralocha and Creadion, on the other. The bill is shorter, but much deeper,
and the jugal bar meets it at a greater angle. The quadrate is relatively much larger,
and is rotated backwards by comparison with the other Callaeidae, so that its otic
process meets the lower jaw more nearly at right angles. The cranium is shortened,
and smaller relative to the orbits ; the jugal is very long relative to the skull. The
lower jaw is deep, and bears strong cristae for the attachment of the adductor
musculature. There is no retroarticular process, and the articular is extremely
shallow at its articulation with the quadrate. The prominent sesamoids at this
articulation were noted by Stonor, and their origin and functions are discussed in
detail by Burton (1973). As in the Huia, the external jugomandibular ligament is
absent. The occipitomandibular ligament is not ossified at all.
The jaw musculature is notable for the highly developed adductor musculature,
and the very small M. depressor by comparison with the other Callaeidae. M.
adductor mandibulae externus is considerably more bulky in actual (as well as rela-
tive) size than in Heteralocha, with increased number of fibres and greater com-
plexity. Its aponeuroses are generally stouter, and more branched and subdivided
internally (Fig. 18), providing extra surface for fibre attachment, and consequently
a greater use of pinnate structure. M. pseudotemporalis superficialis is also much
larger than in the Huia and Saddleback, and in dorsal view completely conceals M.
pseudotemporalis profundus, which is much reduced - a consequence of the back-
ward displacement of the quadrate. M. pterygoideus is bulky and the retractor
palatini slip is especially well developed, and prominent in dorsal view. M. depressor
mandibulae is not only smaller than in the other Callaeidae, but structurally simpler,
and entirely lacks a pars anterior. It is unique, however, for the pulley arrangement
between its aponeurosis and the internal jugo-mandibular ligament (Burton, 1973).
The tongue is much broader relative to its length than in the Huia and Saddleback ;
its shape is almost rectangular, and the tip brush-like. The paraglossa which sup-
port it are widely separated. There is no median aponeurosis and M. ceratoglossus
anterior is absent ; otherwise, the hyoid musculature resembles that of the other
two genera of Callaeidae. The palate is provided with horny papillae similar to
those of the other Callaeidae, but the horny lining of both jaws is developed near the
bill tip into raised, papillate bosses. There is a pair of Gil. angularis oris, but a Gl.
palatinae has not been found.
Neck muscles showing variation in their points of attachment in Heteralocha or
other passerines were examined in Callaeas. The two specimens dissected had the
same number and sites of attachment points for the muscles examined. Slips or
muscle components absent in some Huia specimens, but present in the Kokako
specimens, were the insertions of M. spinalis cervicis on 5 and 6 ; M. splenius colli
arising from 7 ; M. intercristalis from 5 to 3 ; the slip of M. ascendens from 8 to 4 ;
and the slip of M. flexor colli brevis to 6. No Mm. pygmaei arise from 12, though
this site is occupied in all but one male Huia specimen. However, an additional M.
36 P. J. K. BURTON
pygmaeus from 7 to 6 is present in the Kokako specimens, though absent in all the
Huias dissected. The Kokakos lacked the slip of M. flexor colli profundus from 6
to 2, present in three of the Huias, and the slip of M. complexus to 6, present in all
the Huias. M. flexor colli brevis in the Callaeas specimens lacks the slip to 6 present
in two male Huias, but has a slip to 5 (unlike Creadion and one female Huia specimen) .
IX. FUNCTIONAL ASPECTS
Jaw mechanism
Next to the form and sexual dimorphism of the bill, the most striking features of the
Huia's cranial morphology are the huge M. depressor mandibulae and associated
skull modifications - the prominent occipital crest, providing extra surface for its
origin, and the very long retroarticular process providing increased leverage for the
muscle. Clearly these adaptations must permit the lower jaw to be depressed with
great force. This can only be necessary if it is to be opened against considerable
resistance by external forces, and it seems certain, therefore, that the Huia was
highly specialized for feeding by 'gaping' or 'prying'. This feeding technique con-
sists basically of thrusting the bill into a potentially food-bearing medium (earth,
w r ood, fruit, etc.) and opening it, to widen the hole and so facilitate exploration or
prey extraction.
Gaping is a technique described from birds of several families. Among passerines,
good examples are furnished by some Sturnidae and many Icteridae. Gaping
behaviour and related modifications of M. depressor mandibulae have been studied
in the Icteridae by Beecher (1951). It is interesting to compare the Huia with other
birds specialized for gaping, and some figures for relative length of the retroarticular
process in several examples, mainly of passerines, are given in Table 2. The nearest
approach is shown by Cacicus solitarius in which the retroarticular process is slightly
longer relative to the lower jaw than in the Huia. However, Cacicus solitarius is a
shorter billed bird ; relative to skull length, its retroarticular process is shorter than
in Heteralocha, a difference which would be even greater but for the Huia's elongated
skull with enlarged occipital crest. It seems clear that the Huia was very highly
specialized for feeding by gaping, perhaps more so than any bird now living.
Buller's account of the Huia's feeding behaviour is an excellent one, but clearly
gives an incomplete picture of the male's excavation methods. Buller remarks that
the captive male used its bill to 'chisel out' pieces of decayed wood in a woodpecker-
like manner. Almost certainly its precise technique must usually have been to
drive the bill into the wood and 'gape' to split portions off ; this is actually quite
different in principle from the methods of woodpeckers which depend purely on
blows. Buller states that the female he observed fed in a quite different manner,
by probing into relatively hard wood. However, M. depressor mandibulae and the
retroarticular process, though somewhat smaller in the female, are still very large by
comparison with other birds, and her capacity for forceful depression of the lower
jaw must also have been very high. Due to the greater length of the female's bill,
less force could be exerted at its tip than in the male ; and the more flexible nature
of its anterior, purely rhamphothecal, portion, appears rather inefficient for gaping.
ANATOMY OF HEAD AND NECK IN THE HUIA
TABLE 2
Relative length of the retroarticular process in single specimens of various birds showing
gaping adaptations, and in all intact Huia specimens used in this study
37
f
S I
(Si
o .a
K
Upupa epops
Phoeniculus purpureus
Phoeniculus aterrimus
Psarocolius decumanus
Psarocolius wagleri
Cacicus cela
Cacicus leucorhamphus
Cacicus solitarius
Sturnella magna
Amblyrhamphus holosericeus
Creatophora cinerea
Sturnus vulgaris
Sturnus contra
Heteralocha acutirostris, $
(mean of 3)
Heteralocha acutirostris, $
(mean of 2)
Pyrrhocorax pyrrhocorax
81-0
60-4
45'4
65-2
72-6
43'i
48-7
53'5
44'2
45'4
41-7
43'9
49-5
89-4
109-5
77-0
6-2
4'4
7-8
7.9
4-0
5'2
10-3
5'5
8-0
2-8
5'0
4'7
16-4
I3-5
5'4
25'4
26-3
20-5
36-9
37'0
29-1
29-8
31-1
31-3
28-0
28-8
29-4
27-8
46-8
40-9
0-07
o-io
O'lO
O'I2
O'll
O'O9
O'll
0-19
0*12
0-18
0-07
O'll
0-09
0-18
0-07
0'22
0'24
O'2I
0'2I
0'2I
0-14
0-17
o'33
0-17
0-29
O'lO
0-17
0-17
0-31
0-13
Nevertheless, gaping must also have been an important part of her repertoire of
feeding techniques in addition to exploratory probing (as described by Buller) for
which the decurved bill shape is evidently adapted. In probing timber tunnelled
by beetles, it seems feasible that the bill might occasionally be inserted through a
crack into a larger cavity, so that gaping could take place with contact only in the
more rigid basal part of the bill. However, the possibility remains that the female's
apparent gaping adaptations reflect principally the shared genotype of the two sexes
(see under Sexual Dimorphism).
In addition to its large size, M. depressor mandibulae in the Huia also shows
interesting structural complexities. A major contribution to the force of depression
is obviously provided by the large mass arising posteriorly and dorsally on the
cranium, since this is not only bulky, but has the longest moment arm due to its
insertion near the posterior extremity of the retroarticular process. However, the
38 P. J. K. BURTON
anterior parts of the muscle also exhibit various modifications. Several aponeuroses
are present, serving as the basis for pinnate fibre arrangements ; this may be related
to their short working distance, for which pinnate structure should provide more
forceful contraction than parallel fibred muscles of the same physiological cross
section (Gans and Bock, 1965). The pars anterior of the muscle (present also in
C reaction) is of particular interest, since it appears not to have been described pre-
viously in any other bird, and was certainly absent from gaping species of other
families dissected during this study. Its functions are hard to surmise, and a
satisfactory explanation will probably require a much more detailed knowledge of
the jaw mechanics involved in gaping than is available at present.
Except for fibres inserting on the internal process, the greater bulk of M. depressor
mand
