t ai ayare : S Hate Patent: Hitsseuereticneaety BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY VOL IV 1956—1957 PRINTED BY ORDER OF THE TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) LONDON: 1957 DATES OF PUBLICATION OF THE PARTS No. 1 27 March 1956 No. 2. 17 August 1956 No. 3. 17 August 1956 No. 4 28 September 1956 No. 5 23 November 1956 No. 6. 23 November 1956 No. 7. 26 February 1957 No. 8. 6 December 1956 No. 9. 18 January 1957 PRINTED IN GREAT BRITAIN AT THE BARTHOLOMEW PRESS DORKING BY ADLARD AND SON, LTD, CONTENTS ZOOLOGY VOLUME 4 British Mites of the subfamily Macrochelinae Tragardh (Gamasina- Macrochelidae). By G. OWEN EVANS and E. BROWNING (Pls. 1-4) The evolution of Ratites. By SIR GAVIN DE BEER (Pls. 5-9) Studies on the Trichiuroid Fishes—3. A preliminary revision of the family Trichiuridae. By DENYS W. TUCKER (PI. 10) Variation, relationships and evolution in the Pachycephala pectoralis superspecies. By IAN C. J. GALBRAITH A revision of the Lake Victoria Haplochromis species (Pisces, Cichlidae) Part 1: H. obliquidens Hilgend., H. nigricans (Blgr.), H. nuchi- squamulatus (Hilgend.) and H. lividus, sp.n. By P. H. GREENWOOD A systematic revision of the fishes of the Teleost family Carapidae (Percomorphi, Blennioidea), with descriptions of two new species. By D. C. ARNOLD Variation in the western Zosteropidae (Aves). By R. E. MOREAU Sagitta planctonis and related forms. BY P. M. DAVID (Pl. 11) Evolutionary trends in the classification of Capitate hydroids and medusae. By WILLIAM J. REES (Pls. 12-13) Index to Volume 4 57 131 223 INDEX TO VOL. IV The page numbers of the principal references and the new taxonomic names are printed in CLARENDON type. abyssinica, Zosterops ; : 5 nS 4: Acarus . : . . Pt 235 Acaulidae 453-534 Acaulis : 4 466, 481, 509 Acauloidea_ . : : 523 aciculatus, Nothrholaspis 46, 48 aciculatus, Macrocheles (Nothrholaspis) 4c acus, Carapus 2 247-205 Aepyornis . : - 64 aldabrensis, Zostecops maderaspatana = 403 alfurorum, Pachycephala macrorhyncha . 208 allmani, Hydractinia - 476, 478 ambigua, Pachycephala pectoralis 3 . 206 Amphinema . ‘ 4 0 - 481 Andrholaspis 5 A : 5 3 4 anglicus, Coprholaspis 4, 16, 53,54 Annulella 5 5 4 4 : - 506 Anthomedusae cs 3 3 . 520 anzac, Assurger. F 5 fi 2 07 Aphanopodinae 5 “ 3 : 77 Aphanopus_ . di : : : - 81-85 Apteryx a : : 3 5 OS Archaeopteryx 0 4 é : 60-68 arctica, Cyanea 5 4 4 : - 483 arenosa, Aselomaris ci : : = 476 Areolaspinae Q 0 é : é 9 Arum . 0 é . 2 - 467, 509 Aselomaris . : Q ; - 476, 481 Assurger d i 3 9 4 5 ols) Asyncoryne . 2 : . - 459, 509 Asyncorynidae é 0 0 r - 524 Athecata : . 520 atromaculata, Pachycephala pectoralis - 206 aurantiiventris, Pachycephala pectoralis . 198 aurata, Euphysa 460, 466, 486, 492, 505, 507, 519, 521 badius, Holostaspis c Z 16 balim, Pachycephala pectoralis . - 200 banksiana, Pachycephala pectoralis . > 208 bartoni, Pachycephala soror . 5 - 196 bella, Pachycephala pectoralis ; - 198 Benthodesmus 5 : - A 85-90 bermudensis, Carapus_ . : . 270, 272 birpex, Carapus A : : F . 265 blondina, Lizzia_ . . : : me 4k boraborensis, Carapus_. 5 e279 borbonica, Zosterops borbontea : 403 Boreohydra . c 3 ; : + 507 Boreohydrinae 5 0 - 5 - 521 Bougainvillia. z 476, 481 bougainvillei, Pachycephala pectorallis 2) exon Branchiocerianthinae - : - 522 Branchiocerianthus : = . = 472 britannica, Bougainvillia + 481 brunneipectus, Pachycephala pectoralis » 208: buruensis, Pachycephala pectoralis . aE ZO caledonica, Pachycephala pectoralis e200 calliope, Pachycephala pectoralis’ . on Medey? caninus, Carapus . . A . 282 capensis, Myriothela “ = 2 - 468 capillare, Eudendrium 5 7 70) capillata, Cyanea . - 5 - 483 Capitata : 4 j 520 Carapus ; 247 302 carbo, Aphanopus i 83 carinatus, Gamasus : : “i : 25 carinata, Nothrholaspis . : 25 carinatus, Macrocheles 12, 25, Bel 34, "36, 53 carnea, Podocoryne 476, 478 castaneus, Macrocheles . ‘ : é 2 Casuarius A “ A 0 : 63 caudatus, Lepidopus Cc Os centralis, Pachycephala santas a - 198 chlorura, Pachycephala pectoralis . eEZOe christophori, Pachycephala pectoralis > 198) cinereus, Carapus . - - 295 cinnamomea, Pachycephala pectoralis =) BLOF, citreogaster, Pachycephala pectoralis Sze Cladocoryne A : a é SS Gp Cladocorynidae z 5 £ : a cya! Cladonema : 462, 464, 511, 519 Cladonemidae a 6 é Cag 25 Clava . e - 476 clio, Bachycepbala pectoralis 202 cocksi, Arum 467, 487, os. 509 cognatus, Macrocheles_ . 28 collaris, Pachycephala pectoralis c e205 comorensis, Zosterops maderaspatana - 403 conferta, Dicoryne 2 476 contempta, Pachycephala pectoralis 5 lop conybearei, Heterocordyle 3 476 Coprholaspis . 4, 7, 10, 16, "53, 54 Cordylophora : : : 7470, cornucopiae, Merona 7 a 3 - 476 Corymorpha 462-508 Corymorphidae 453-534 Corymorphinae Corynidae Corynoidea costata, Zanclea “474, 499, 512, 519, 526 INDEX 537 521 453-534 524 cristatus, Tentoriceps 110 crocea, Tubularia é 481 cucullata, Pachycephala ee toralis a Tezon curvirostris, Zosterops curvirostris . - 403 cuspis, Carapus 267 Cyanea F 2483 cyclopum, Pachycephala lschlegelii “ LOS Cyrtocheles i 4 Cytaeis 470 dahli, Pachycephala pectoralis . ¢ LGS dammeriana, Pachycephala Bae - 202 decoloratus, Gamasus a 32 decoloratus, Macrocheles 12, 32, 33, 34, 53 Dendrocoryne 475, 513 dentatus, Echiodon 3 ‘i . 292 dentatus, Macrholaspis - 4, 46, 48, 51, 53 dichotoma, Eleutheria 500, 512, 519, 525 Dicoryne : 476 dinema, ‘Amphinema 481 Dinornis 64 Diplospinus 78-81 Dipurena 400, 475, 493, 498 Dissoloncha . 4, 10 domestica, Musca 13 Dromaeus 63 drummondii, Echiodon 288 dubius, Carapus 270 dumortieri, Ectopleura echinata, Hydractinia 491, 496, 509, 519 es 479, 481 Echiodon 288-294 Ectopleura 491, 509 efatensis, Pachycephala pectoralis : « 208 Eleutheria F 500, 512, 519 Eleutheriidae 52 elongatus, Benthodesmus 88 Encheliophis 295-298 Eudendrium . 476 Euphysa 460-507 Euphysinae Sez eurycricota, Zosterops “ - 363 everetti, Pachycephala pectoralis 5 De 2OG' eximia, Sarsia 6 519 Eupleurogrammus . 102 Evoxymetopon 97 farcta, Euphysa a . 486 feminina, Pachycephala pectoralis f 2 198 fergussonis, Pachycephala pectoralis a n207, filiformis, Staurocoryne . 462 fimicola, Nothrholaspis 36, 38 flavifrons, Pachycephala 136-222 floccosa, Cladocoryne 511, 524 Formica " 5 13 fulginosa, Pachycephala pectoralis : 7) zoo fulviventris, Pachycephala pectoralis = rQ6! fulvotincta, Pachycephala pectoralis gb: fuscoflava, Pachycephala pectoralis . 202 fusiformis, Rhizogeton : 476, 478 Gamasina-Veigaiaidae . 7, Gamasus 18, 16, 25, 30, 32, 42, Gy, 48 Garveia 4 : 5 «476 Geholaspis - 4-10, 41-43, 53 gemmifera, Sarsia . 493, 519 Gempylina 21-127 gigantea, Monocoryne 488, 509, 523 gilolonis, Pachycephala pectoralis . zoo) glaber, Macrocheles 45/10), 00, 16, 57.00. 53 glabra, Holostaspis a : 16 glacialis, Corymorpha_. ‘ ~ 496 gladiator, Macrocheles 36, 38, 53 glaucura, Pachycephala pectoralis_ . . 200 gloriosus Macrocheles (Nothrholaspis) = es goodsoni, Pachycephala aa 5 5. Os} Gotoea 5 - - 493 gracilis, Diplospinus : , af) se gracilis, Encheliophis (Jordanicus) ‘ . 299 gracilis, Solanderia 5 = 526 graefi, Pachycephala peatoralie . 7 198 groenlandica, Corymorpha : 0 - 406 haeckeli, Margelopsis : 482, 508, 522 haesitata, Zosterops curvirosttis t a 440)3} Halocordyle . - b . 460, 474, 510 Halocordylidae ¢ : : : 524 halterata, Dipurena é : ; 47s, 493 hancocki, Encheliophis_ . ‘ ‘ . 298 Haplochromis ci : j + 225-243 Hesperornis . : 2 : LOY Heterocordyle a f c - 476 Holostaspella bs 7,9, 10, 51-53 Holostaspis - 13, 27, 38, 44, 48, 51 Holostaspis (Holostaspella) . 3 51 Holostaspis Pisentii r : - : 13 Holaspulus . : : : A 2 9 homei, Carapus ‘6 3 £ 1273 hooperi, Stylactis . : ¢ : +) 480 houlti, Carapus : “ c : 6 atk) hulli, Macrocheles . : o , a 53 hulli, Nothrholaspis : : 3 Bes Hybocodon . : : : 491 Hydractinia . 475, 476, 481 Hydrocoryne : : é 500, 512 Hydrocorynidae . 7 3 - 525 hypochthonius, Macrocheles é j a 25 Hypolytus . fs 3 é . 460, 506 Ichthyornis 7 5 : : 67 imperator, Branchiocerianthus ° Bal ye indivisa, Tubularia. o c - 496, 522 inkermanika, Ostroumovia 6 A etd 83 538 INDEX insignitus, Macrocheles 11, 24, 53 intermedius, Eupleurogrammus Fs =) Mog japonica, Cytaeis . ¢ - 476 javana, Pachycephala pectoralis - 196 jordani, Encheliophis 298 Jordanicus . c - 299 jubilarii, Pachycephala pectoralis 5 = 200 kagoshimanus, Carapus . . 278 kandavensis, Pachycephala pectoralis - 204 kirki, Zosterops senegalensis . . + 403 klossi, Pachycephala soror 5 . - 196 lacustris, Cordylophora 470 lamarcki, Cyanea 4 b . 483 larynx, Tubularia . : 4 5 - 496 lateralis, Zosterops : - 316 lauana, Pachycephala pectoralis . . 204 Lepidopodinae : r : : - 89 Lepidopus a ° 90-97 Lepturacanthus” . 3 m 4 = EXO lepturus, Trichiurus : E : a 4! Leuckartiara : 2 3 - 476 linearis, Bougainvillia. 5 - 476 littayei, Pachycephala pectoralis 5 2 20% lividus, Haplochromis : 3 . 232 Lizzia . : : 3 5 . . 481 Longicheles : 4, 42, 44 longispinosus, Gamasus . : Az longispinosus, Geholaspis, (Geholaspis) - 42 longispinosus, Holostaspis : “ : 2 longulus, Geholaspis (Longicheles) . . 44 longulus, Holostaspis —. 2 i caer Macrholaspis 4,5, 7, 10, 46-50 Macrocheles . 5 + 4-53 Macrocheles (Coprholaspis) 2 . ena: Macrocheles (Monoplites) 16, 41 Macrocheles (Nothrholaspis) 41, 48 Macrochelidae . 4,9, 53 Macrochelinae : 45.9) 50, 53 macrorhyncha, Pachyceukale pectoralis = 202 maderaspatana, Zosterops maderaspatana 403 Malacirops 309-433 mandibularis, Geholaspis fanereneles) 44, 45, 47, 53 mandibularis, Holostaspis a aa margaritiferae, Carapus (Onuxodon). 285 Margelopsidae 5 453-534 Margelopsinae : . 5 5 522 Margelopsis : 5 - 482, 508 marginatus, Acarus fs 11, 38 marginatus, Holostaspis . 19, 36, 38 marginatus, Macrocheles 4, 16 marri, Sagitta 443 matrius, Macrocheles 12, 82, 33, 30 matrius, Nothrholaspis_ . 4 a 34 mauritiana, Zosterops borbonica : - 403 mayottensis, Zosterops semiflava . - 403 mediterranea, Porpita . - + 519 melanonota, Pachycephala pectoralis <4 007 melanops, Pachycephala pectoralis . 204 melanoptera, Pachycephala pectoralis 198 melanura, Pachycephala pectoralis . 199 mentalis, Pachycephala pectoralis 197 merdarius, Holostaspis 21 merdarius, Macrocheles . 11, 21, 22 Merona 476 merula, Saxicola 209 michaeli, Aselomaris 481 minimus, Macrocheles 44, 53 mirabilis, Pelagohydra_. c 470, 508, 522 misakinensis, Dendrocoryne = Sus misimae, Pachycephala pectoralis 209 miurensis, Hydrocoryne . 3 500, 512, 525 modesta, Zosterops + 403 Monocoryne . 488, 509 Monocoryninae - 523 Monoplites 3 10, 16 montana, Nothrholaspis 4 23 montanus, Macrocheles 23, 24, 5 53 mouroniensis, Zosterops . 403 multistriatus, Diplospinus 79 Musca . Q 13 minseaedomesticac) ‘Macrocheles 6-12, 13 muscoides, Bougainvillia. 476 muticus, Eupleurogrammus 105 myersi, Cladonema. - 464 Myriothela 467; 468, 487 Myriothelidae 453-534 Myriothelinae 523 nemoralis, Macrocheles (Nothrholaspis) 41 Neoholaspis . 5 4 nigricans, Haplochromis 237 Nothrholaspis ‘ 10, 40, 53 nuchisquamulatus, Haplochromis z 241 nutans, Corymorpha toe 508, 519, 521 nutans, Garveia 470 nutricula, Turritopsis 78: 478 Obelia . 488 obiensis, Pachycephala pectoralis 197 obliquidens, Haplochromis 226 obscurior, Pachycephala schlegelii 195 obvoluta, Hypolytus ‘i 460 occidentalis Macrocheles (Notnrholaspis) 28 octona, Leuckartiara 476 octopunctata, Rathkea 481 Onuxodon 284 opacus, Gamasus 46, 48 opacus, Macrholaspis 4, 48 ophiogaster, Dipurena c 493 orioloides, Pachycephala pectoralis a ey ornata, Holostaspella é 51, 52, 53 ornatus, Holostaspis = 5 ornata, Pachycephala pectoralis 204 Ornitholestes ci 2 6a Ostroumovia 483, 506 ottomeyeri, Pachycephala pectoralis Bh 3 INDEX 539 oudemansii, Macrocheles (Monoplites) 4, 16, 36 owasianus, Carapus : 3 : . 278 Pachycephala i 131-222 pallida, Zosterops . : eet palustris, Macrocheles (Monoplites) : . 41 pannosus, Macrocheles (Nothrholaspis) . 414 Parholaspis . F : 9 parmulatus, Macrocheles ‘(Nothrholaspis) . 4 parvibrachium, Carapus (Onuxodon) . 284 parvipinnus, Carapus’. 0 a . 279 pectoralis, Pachycephala. 4 131-222 pectoralis, Pachycephala pectoralis . . 200 Pelagohydra . - c 5 2 - 508 Pelagohydrinae . ee 22 pelengensis, Pachycephala pectoralis) on 425) penicilliger, Macrocheles 12, 27, 41, 53 penola, Myriothela : 3 F . 467 peregrinus, Hypolytus. 460, 506 phrygia, Myriothela 487, 523 pindae, Carapus . 3 : “ a cate pisentii, Macrocheles 5 . ~ it, Ao 04! planctonis, Sagitta 437-540 plumipes, Macrocheles 36, 38, 53 plumiventris, Macrocheles 12, 36-38, 41, 53 Podocoryne . 2 : ° . eAy70) poeyi, Evoxymetopon . : Fi ~ fOr Porpita. - 5 3 = ES) primarius, Acaulis . 466, 481, 509, 523 producta, Stauridiosarsia 463, 498, 510 prolifer, Hybocodon 3 . 491, 496, 505 prolifera, Sarsia 493, 519 Protoholaspinae . : 4 ‘4 " 9 Protoholaspis : 2 : : A 9 Ptilocodiidae 3 A { « 525 Ptilocodium 474) 503 punetoscutatus, Macrocheles 11, 18, 19, 53 pusilla, Coryne 498, 525 queenslandica, Pachycephala pectoralis . 200 radiatum, Cladonema 460, 511, 519, 525 ramosum, Eudendrium : : - 476 Rathkea 5 : 4 - “ 4cr reedi, Carapus i 5 : 4 . 287 regalis, Tubularia . : 2 5 - 496 rendahli, Carapus . é F 3 . 294 rendovae, Zosterops 0 , : Cee repens, Ptilocodium : 513, 525 Rhea . x = : c . 63, 64 Rhizogeton . , : . 476 Rhizorhagium 0 : : : . 476 Riparia 2 3 : : 6 30 riparia, Riparia. F : ~ 36 Rosalinda. 0 : 0 - 513 roseum, Rhizorhagium r : 2 - 476 rothamstedensis, Macrocheles é 11, 15, 53 rufa, Formica ' A 3 13 tyniensis, Asyncoryne 459, 509, 524 sagamianus, a ala (Jordanicus) . 301 Sagitta : 437-450 salomonis, Paebyeernalel - : e207, sanfordi, Pachycephala a aan 7 Ge ey} Sarsia . 5 - 493, 519 savala, Lepturacanthus 4 é 3 DLO) scaber, Trox, é c 5 5 AS) Scarabaeus . ¢ - 4 ° 13 schlegelii, Pachyeephale a : 136-222 sculpta, Holostaspis (Holostaspella) sl) WS semiflava, Zosterops semiflava é . 403 semipunctatus, Scarabaeus”. : : 13 senegalensis, Zosterops . : = 3nd sharpei, Pachycephala pectoralis ‘ "202 simonyi, Benthodesmus c 5 88 simplex, Boreohydra xo, 519, 522 singularis, Tricyclusa 462-523 Solanderiidae 6 5 ‘ é 7520) soror, Pachycephala : Q : 136-222 Speirops : : 3090-433 Sphaerocerus : é 53 spinicauda, Pachycephala pectoralis - 199 squamata, Clava. “ - 476 Stauridiosarsia 460, Kens 498, 510 Staurocoryne. 3 5 = 5 - 462 stercorarius,Gamasus_ . q M G 16 stercorarius, Geotrupes ¢ 5 18 strangulata, Dipurena_ . 493, 498 Struthio : c , 9 5 (% subbadius, Holostaspis 0 19 subbadius, Macrocheles Il, 19, 34, 36, 53 submotus, Macrocheles . II, 12, 28, 29, 53 superbus, Macrocheles 9, 12, 38, 39 superciliaris, Boupainvillia : é . 481 tabarensis, Pachycephala pectoralis . oye} taeniatus, Evoxymetopon : F : 99 tardior, Macrocheles (Monoplites) . - 41 tardus, Gamasus_ . : : 13, 30 tardus, Macrocheles 12, 30, 31, 53 tenuipes, Holaspulus : é : 9 Tentoriceps . A : , 2 sO. tenuis, Brenined esanne i 5 : 3 89 terreus, Holostaspis ¢ 7 48 teysmanni, eae pectoralis | : 206 Thyrsitina . ¢ a 5 ited tiarella, Halocordyle 3 510, 524 tidorensis, Pachycephala pectoralis 2 = 007 torquata, Pachycephala a 3 =. 198 Trichiuridae . fs 0 . 73-130 Trichiurinae . é : : : SEZ Trichiurus . d : : Pe | Tricyclusa 462, 482, 480, 508 Tricyclusidae . « , 453-534 Tricyclusoidea 5 - . we522 tridentinus, Miecrocheles!: ; : a 30 Tringa . z : - : v 316% Drox, 4 c 2 ° 28 Tubularia 470; 481, 496 Tubulariidae 453-534 540 INDEX Tubularoidea : : " - 520 virens, Zosterops . : - 314 tubulosa, Sarsia. cj ; 403, 498, 505 viridipectus, Pachycephala schlegelii Be 742h5) Turritopsis . : : dt 2 . 476 vitiensis, Pachycephala pectoralis. = Ow typica, Gotoea é 4 Fi - 493 véltzkowi, Zosterops maderaspatana - 403 urceolus, Branchiocerianthus . - 472, 522 whitneyi, Pachycephala esac F - 199 utupuae, Pachycephala pectoralis_ . . 204 williami, Rosalinda : 2 “ 513~ vagabundus, Macrocheles : 3 51 xanthoprocta, Pachycephala pectoralis . 201 vanikorensis, Eee nal pectoralis 5 202 xantusi, Lepidopus . . 5 - 95 Veigaia 5 : 5 : 0 7 velella, Velella : ; : ‘ - 519 Zanclea . : . 474, 499, 512, 519 vermicularis, Encheliophis ; : . 295 Zancleidae . 5 F E 1520 veterrimus, Macrocheles . 4 4 2 6 zetesios, Sagitta . : ‘ - 437, 443 violetae, Pachycephala pectoralis. ae, Zosterops.. . 5 : + 3090-433 fo 4 , 23 ae re Pais ESE SBC) _ G OWEN EVANS _E. BROWNING. BRITISH MITES OF THE SUBFAMILY MACROCHELINAE TRAGARDH (GAMASINA—MACROCHELIDAE) as BY G. OWEN EVANS AND E. BROWNING Pp. 1-55; Pls. 1-4; 85 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 4 No. 1 LONDON : 1956 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), «stituted in 1949, is issued in five series corresponding to the Departments of the Museum, and an Historical Series. Parts appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be compiled within one calendar year. This paper is Vol. 4, No. i of the Zoological series. PRINTED BY ORDER OF THE TRUSTEES OF THE BRITISH MUSEUM Issued March, 1956. Price Seventeen Shillings and Sixpence. BRITISH MITES “OF THE SUBFAMILY MACROCHELINAE TRAGARDH (GAMASINA—MACROCHELIDAE) By G. OWEN EVANS anv E. BROWNING CONTENTS Page INTRODUCTION . : : B : : : 4 4 EXTERNAL MorpHotocy 2 : : c 6 5 ° c 4 CLASSIFICATION . a 0 : é 3 : "3 : 9 Genus Macrocheles Latr. 4 : ° 2 : 5 as) Macrocheles muscaedomesticae (Scopoli) 0 5 0 ¢ : An) 173 Macrocheles pisentii (Berl.) . 2 6 : 5 ¢ 5 5 13 Macrocheles vothamstedensis sp.nov. . : 9 : é at ype Macrocheles glabey (Miiller) . ss : : é 3 5 ue Macrocheles punctoscutatus sp. nov. é 5 : 5 : ELS Macrocheles subbadius (Berl.) : 5 : c : “ 0 19 Macrocheles insignitus Berl. c c a 2 0 : at Macrocheles merdarius (Berl.) , : : : : : 0 | eB Macrocheles montanus (Willmann) : : : 2 2 eR Macrocheles carinatus (C. L. Koch) 3 “ : : GAB Macrocheles penicilligey (Berl.) . é q : : : a 27; Macrocheles submotus Falconer. 5 : a : = 2 Macrocheles tardus (C.L. Koch) . 5 : ‘ é 5 - 30 Macrocheles decolovatus (C.L. Koch) . : 6 5 5 ese Macrocheles matrius Hull. : ‘i a 5 n é a Si Macrocheles plumiventris Hull 0 : a é : : 30 Macrocheles superbus Hull . : 6 ; a 4 . so Species incertae sedis . ; ; a : : c : oxi Genus Geholaspis Berl. s. lat. : : : : eA Geholaspis (Geholaspis) longispinosus (Kramer) : ; j 2 6B Geholaspis (Longicheles) longulus (Berl.) : : é J a AML Geholaspis (Longicheles) mandibularis (Berl.) . : 4 : - 44 Genus Macrholaspis Oudemans_ A F 6 4 , 2 tS Macrholaspis opacus (C. L. Koch) 5 . : . : . 48 Macrholaspis dentatus sp.nov. . é 5 : A : - 48 Genus Holostaspella Berl. . 5 : < : . 5 Bi Holostaspella ornata (Berl.) . : o : : : : ne ah SUMMARY co : : - 5 : : : : : den abis} ACKNOWLEDGMENTS. : 5 , : : . 3 c - 54 REFERENCES 5 3 ; : 6 : : : i : 54 SYNOPSIS The classification of the Macrochelidae is discussed, with particular reference to the British species, and three new species are described and figured. ZOOL, 4, I. 1 4 BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH INTRODUCTION THE recent lists of the genera of the Macrochelidae follow the classification proposed by Berlese (1918), the only comprehensive work on the family. Vitzthum (1941) recognizes eight genera and eleven subgenera to which must be added the genera Neoholaspis Turk, 1948 (? syn: Macrocheles Latr. s. str.) and Andrholaspis Turk, 1948, and the subgenera Cyrtocheles Valle, 1953 and Longicheles Valle, 1953 of Geholaspis Berl. s. lat. The British Macrochelidae have been investigated by Hull (1918 and 1925), Falconer (1923 and 1924) and Turk (1946). In 1918, Hull keyed ten species of Macrocheles, of which four were considered to benew. Later, in 1925, he described ten more new species and proposed a new name (Macrocheles (Monoplites) oudemansit) for Macrocheles marginatus Oudemans, 1901 nec Hermann, 1804. He used the chaeto- taxy of the dorsal shield and the form of the ventral shields as his chief taxonomic criteria and although these characters have been proved subsequently to be useful key characters, his descriptions and figures are so inadequate (and often inaccurate) that a number of the species cannot be recognised with certainty. Falconer’s contribution to the study of the family consists of descriptions and figures of two new species and the erection of a new subgenus (Dissoloncha) of Macrocheles. Turk (1946) has described one new species, Coprholaspis anglicus, from under wet wood at Reskadinnick, Cornwall, and has keyed British species of the genus Coprho- laspis Berl. The object of the present work is to redescribe and figure the known British species of the Macrochelinae. We have followed Sellnick’s interpretation of the species of C. L. Koch and Scopoli (Sellnick, 1931 and 1940). EXTERNAL MORPHOLOGY The following account of the external morphology of the British species of the Macrochelinae refers to the adult stages only and will serve as a general introduction to the characters of taxonomic importance used in the keys to species. Dorsal shield : The dorsal shield in all British species is entire and covers practically the whole of the dorsum of the mite. The ornamentation of the shield shows con- siderable variety in form, for example, the shield may be faintly reticulated as in Macrocheles glaber (Miller), strongly reticulated and punctate as in Macrocheles tardus (C. L. Koch) or densely covered with minute tubercles as in Geholaspis (Longicheles) mandibularis (Berl.). Its lateral margin may be smooth or serrated. The number of setae on the shield is remarkably constant, twenty-eight pairs in the female, except in three species (Macrocheles montana (Willm.), Macrholaspis opacus (C. L. Koch) and Macrholaspis dentatus sp.n.). The chaetotactic pattern and structure of the setae (whether simple, serrated or plumose) afford valuable taxonomic criteria. Sellnick (1942) and Valle (1955) have already used the chaetotaxy of the shield in their works on Macrocheles and Geholaspis s. lat., respectively. The nomenclature used for the dorsal chaetotaxy in the present work is given in Text-fig. 5. The twenty- th eS 4 BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH 5 eight pairs of setae of the female are divided into four longitudinal rows; a dorsal series (D) of eight pairs, a median series (M) of four pairs, a lateral series (L) of six pairs and a marginal series (Mg) of ten pairs. This division is purely artificial and is not based on the post-embryonic developmental sequence. In the males, two groups Fics. 1-4. The chaetotaxy of the ventri-anal shield in females of Macrholaspis Oude- mans (Fig. 1), Macrocheles Latr. (Fig. 2), Holostaspella Berl. (Fig. 3), and Geholaspis Berl. (Fig. 4), a, anterior pre-anal seta ; b, median pre-anal seta ; c, posterior pre-anal seta; d, antero-lateral pre-anal seta ; e, postero-lateral pre-anal seta. can be recognized, namely, those which have the same chaetotactic pattern as the female and those which have a greater number of setae on the shield. In the latter, setae are added to the shield through its extension laterally to incorporate a number of the setae (the extra-marginal setae) normally situated on the lateral interscutal membrane. The anterior margin of the shield in the genera Macrocheles, Macrholaspis and Geholaspis is gently rounded with the vertical setae (Dz) situated medially at a short 6 BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH Fic. 5. Chaeto se aS of the dorsal shield in Beet cae muscaedomesticae (Scopoli), female. D1— do: ene series; M1—M4, median series; L1—L6, lateral series ; Mgr-Mgro, ma: ate al si BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH 7 distance from the margin. In Holostaspella, however, the vertical setae are no longer on the summit of the shield but on an outgrowth from it (Text-fig. 82). In addition to the setae, the shield is provided with a number of pore-like structures. There are normally twenty-two pairs of these “‘ pores’. Tritosternum. This structure is well-developed throughout the group and consists of a rectangular base, longer than wide, and a pair of long, pilose lacinae. Ventral shields. In the female, the ventral shields comprise the sternal, metaster- nals, genital, ventri-anal and metapodals. The sternal shield is strongly sclerotized and carries three pairs of setae (hr, h2 and h3) and two pairs of pores. The ornamentation of the shield is usually character- istic of a species or a group of species. Berlese (1918) used the ornamentation of the shield as a major character for subdividing the subgenus Coprholaspis and introduced a system of nomenclature for the lines and punctate areas forming the basic pattern (Text-fig. 2). The writers have referred, in the main, to a photograph of the sternal shield instead of attempting a description of the ornamentation. The metasternal shields are paired and free, i.e. they are not fused with the sternal or endopodal shields. Each bears a seta and usually a “‘ pore”’. The genital shield lies between coxae IV and carries a pair of genital setae. It is invariably reticulated and punctured, and is provided with accessory sclerites as in the genus Vergaia Oudms. (Gamasina—Veigaiaidae). The lateral sclerites are usually well-developed, but the median sclerite is often only weakly sclerotized. The region posterior to coxae IV is largely occupied by a ventri-anal shield bearing two (Macrholaspis), three (Macrocheles), four (Holostaspella) or five pairs (Geholaspis) of pre-anal setae. The nomenclature for these setae is given in Text-figs. 3-6. In addition to the pre-anals the shield bears the normal three setae associated with the anus, namely, the paired para-anals and the post-anal seta. The surface of the shield is usually reticulated and punctured. The interscutal membrane between the genital and ventri-anal shields may be provided with sclerotised platelets. The metapodal shields are relatively small and inconspicuous. In the male, the ventral surface may be covered by a sterniti-genital shield in the region of coxae I-IV and a separate ventri-anal shield posterior to coxae IV or, by a holoventral shield, i.e. a fused sterniti-genital and ventri-anal shield. The genital orifice is prae-sternal. Stigmata, peritremes and peritrematal shields. The stigmata, one on each side of the idiosoma, are situated between coxae III and IV. The peritremes are well- developed and extend beyond the level of coxae I. Each forms a U-shaped loop in the region of the stigma (Text-fig. 6). The peritrematal shield is fused anteriorly with the dorsal shield, but is free in its proximal half. Gnathosoma. The gnathosoma is typical of that found in the free-living Gamasina. Ventrally, it is provided with four pairs of setae and a distinct ventral or capitular groove (Text-fig. 7). The latter has five to seven transverse rows of denticles. The corniculi are well-developed and the internal malae long and pilose. The salivary styli are also prominent. The five free segments of the pedipalp bear simple, rod-like or spatulate setae. The chaetotactic formula for the trochanter, femur and genu is 8 BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH Fics. 6-9. Macrocheles muscaedomesticae (Scopoli), female. Fig. 6, lateral view. Fig. 7, venter of gnathosoma. Fig. 8, tectum. Fig. 9, chelicera. Abbreviations: c.s., capitular seta ; d.sh., dorsal shield ; ext, mg., extra-marginal series ; ext.p.y.; external posterior rostrals; int.p.y.,, internal posterior rostrals; per., peritreme ; per.sh., peritrematal shield; r., rostral seta; st¢., stigma, BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH 9 2-5-6. The specialized seta on the inner basal angle of the palptarsus is three pronged. This segment also bears a conspicuous long, upright, rod-like seta distally. The tectum is extremely variable in form. In the majority of the species it consists of three anteriorly directed processes. These may be free (Text-fig. 8) or each lateral process partly fused with the median (Text-fig. 31). The lateral processes may be smooth and entire or divided and serrated. The median process is usually strongly setose and bifurcate distally. In Macrocheles superbus Hull and Geholaspis s. str. the tectum is simply produced into a single anteriorly directed process, variously divided distally (Text-figs. 57 and 64). The chelicerae in both sexes are chelate-dentate. The teeth are massive and ridged, or conical and smooth. The pilus dentilis, dorsal seta and pore, lyriform pore and ventral setae are well-developed. In the male the movable digit is provided with a strong spermatophoral process. This may be long and slender or short and inflated (Text-figs. 39 and 60). Ambulatory appendages. All the legs are six-segmented with the ultimate segment incompletely divided into a metatarsus and tarsus. Leg I is without an ambulacral apparatus (pulvillus and claws) and terminates in a number of sensory setae. This is a characteristic feature of the family. Legs II-IV, however, have well-developed lobate pulvilli and claws. In the females, spurs are present on the femur and tarsus of leg II in the genus Holostaspella only (Text-fig. 85). The femur, genu and tibia of Leg II and often one or more segments of leg IV are spurred in the males. CLASSIFICATION Tragardh (1952) divided the Macrochelidae into three subfamilies as follows : “ 1. Metasternal shields connected with the sternal shield through a narrow bridge Protoholaspinae Tragardh, 1949 —. Metasternal shields free . : ; Z ; : , 5 5 A : Fa . Peritrematic shields not fused with the exopodal shields Macrochelinae Tragardh, 1949 —. Peritrematic shields fused with the exopodal shields . . Aveolaspinae nov. subfam.” The genus Protoholaspis Tragardh 1949, the only member of the Protoholaspinae, is not a typical Macrochelid in that leg I is provided with an ambulacrum and the chelicera lacks the characteristic ventral seta. Further, Tragardh was unable to see the structure of the specialized seta on the inner basal angle of the palptarsus, so that the exact systematic position of the genus must remain in doubt pending the re- examination of the type. The family should, therefore, be considered to consist of the two subfamilies separated in couplet 2 of the above key. The majority of the British species belong to the Macrochelinae ; the Aveolaspinae being represented by Holaspulus tenuipes Berl. and Parholaspis sp., which are intro- duced species that have become established in the Aroid House, Royal Botanic Gardens, Kew. The following separation of the four British genera of the Macro- chelinae is based on Evans (1956) : 10 BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH Key to genera Females 1. Femur of leg II armed with strong spurs ; vertical setae situated on an outgrowth of the dorsal shield ; ventri-anal shield with four pairs of pre-analsetae Holostaspella Berl. —, Femur of leg II unarmed ; vertical setae on the summit of the dorsal shield ; ventri- anal shield with 2, 3 or 5 pairs of pre-anal setae : 5 a 3 : 2. 2. Ventri-anal shield with two pairs of pre-anal setae é : . Macrholapis Oudemans —. Ventri-anal shield with more than two pairs of pre-anal setae 5 6 > 3h 3. Ventri-anal shield with three pairs of pre-anal setae. é . Macrocheles Latreille —. Ventri-anal shield with five pairs of pre-anal setae ; 5 . Geholaspis Berl. s. lat. Genus MACROCHELES Latreille Macrocheles Latreille, P. (1829). In Cuvier, Régne anim., Ed. 2, 4: 282. Coprholaspis Berlese, A. (1918). Redia, 13: 146. Nothrholaspis Berlese, A. (1918). Redia, 13 : 169. Dissoloncha Falconer, W. (1923). Naturalist, Lond. : 151. Monoplites Hull, J. E. (1925). Ann. Mag. nat. Hist. (9), 15: 215. Berlese (1918) divided the genus Macrocheles into four subgenera, namely, Macro- cheles s. str., Coprholaspis, Nothrholaspis and Geholaspis. Sellnick (1940) has shown conclusively that Coprholaspis is a synonym of Macrocheles s. str. and, within recent years, Geholaspis has been given generic status. The remaining two subgenera have been separated, chiefly, by the ornamentation of the sternal shield in the female. In the majority of the species of Macrocheles s. str., this shield is ornamented with distinct lines or punctate lines whereas in Nothrholaspis the ornamentation takes the form of a network of ridges (cf. Pl. 1, fig. 3 and Pl. 3, fig. 15). This distinction is not as definite as supposed by Berlese and, as is often the case with a general character of this nature, an intermediate group, containing species which could belong to either subgenus, is apparent. In view of this, and until a more comprehensive study can be made of both sexes of the known Macrocheles species, especially those in the Berlese Collection, the writers feel it advisible to combine the subgenus Nothrholaspis with Macrocheles. Ecologically, the British species of this genus may be divided into two groups : those living in dung and those inhabiting leaf litter or mosses. The former contains a number of phoretic species, e.g. M. muscaedomesticae and Macrocheles glaber (Miiller), in which the shape and chaetotaxy of the dorsal shield in the male differs considerably from that in the female. The genus Macrocheles may be defined as follows: Dorsal shield in both sexes entire with usually twenty-eight! pairs of setae and twenty-two pairs of “ pores.” Sternal shield in the female with three pairs of setae ; metasternal shields free. Genital shield with a pair of setae and accessory sclerites. Ventri-anal shield with three pairs of pre-anal setae in addition to the three setae normally associated with the anus. Male with sterniti-geniti-ventral shield and separate ventri-anal or with holoventral 1 Macrocheles montanus (Willmann) has twenty-nine to thirty-one pairs of setae on the dorsal shield, and in some males the number of setae may be increased through the lateral extension of the shield to incorporate a number of the extra-marginal setae, BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH II shield. Genital orifice prae-sternal. Hypostome with well-developed corniculi and salivary styli. Pedipalps with five free segments ; chaetotaxy of trochanter, femur and genu being (2-5-6). Specialized seta on the palptarsus with three prongs. Chelicerae chelate dentate with a well developed brush of setae ventro-laterally ; movable digit in the male with a strong spermatophoral process. Legs I without ambulacra ; legs II and often legs IV, spurred in the male. Tyre: Acarus marginatus Hermann, 1804 (= Acarus muscae domesticae Scopoli, 1772). Key to the Females of the British Species of Macrocheles Latr. I. Setae L3 to L5 and Mg3 to Mg6 simple 2. —. Setae L3 to L5 and Mg3 to Mgé serrate, plumose or pencillatel 8. 2. Setae D8 simple or pilose é . c ‘ Be —. Setae D8 short, comb-like (Text-fig. 11) ; sternal shield weakly ornamented ; ventri- anal shield considerably longer than broad ; fixed digit of the chelicera with a row of 5 or 6 small teeth distally (Text-fig. 13) i 5 . Macrocheles pisentii (Berl.) 3. Vertical setae (D1) stout and plumose or long and setiform (Text-fig. 22). Dorsal shield more than 6004 in length. : 5 ¢ 5 : A : 4. —. Vertical setae short spine-like (Text-fig. 24). Dorsal shield less than 500 in length : ; 9 : 2 6 : 2 7. 4. Vertical setae short and Sout, plumose dieaits = setae D4, L1, Lz, L6 and Mgr simple (Text-fig. 17); tibia I approximately equal in length to tarsus (1 : I-o— I-05) .- 5: —. Vertical setae ‘Jong setiform (Text= he, I ay att Snip aaioconrants serrations distally ; ; sternal shield densely punctured; setae D4, Li, Lz, L6 and Mgr pencillate ; tibia I shorter than tarsus I (1 : 1-2-1-44) 0 Macrocheles vothamstedensis sp. nov. 5. Sternal shield with deeply incised transverse median and posterior oblique lines (Pl. 1, fig. 3) ; dorsal setae, except D1, D4 and Lz, simple Macrocheles glabery (Miiller) —. Sternal shield punctured, transverse median and posterior oblique lines poorly developed or absent ; setae D4 simple ¢ ¢ c c : : 6. Ventral shields densely punctured (Pl. 1, fig. 4); setae Mg2, Mg7, Mgg and D8 distinctly serrated (Text-fig. 21) . Macrocheles punctoscutatus sp. nov. —. Sternal shield with fewer large punctures (Pl. i fe, a) setae D1, D8 and L2 only serrate or pencillate . ¢. Macrocheles subbadius (Berl.) 7. Sternal shield ornamented with large ‘punctures (Pl. lp ‘fe. 6) Macrocheles insignitus Berl. —. Sternal shield weakly ornamented with punctate lines (PI. 2, fig. 7) Macrocheles merdarius (Berl.) 8. Dorsal shield with 29% or more pairs of setae (Text-fig. 27) ; median pre-anal seta on or slightly off the line connecting the anterior and posterior pre-anals (PI. 2, fig. 8) 5 ; 5 2 . Macrocheles montanus a ane -. Dorsal shield with 28 pairs of setae 5 2 : 9. g. Vertical setae situated in close proximity to each other, so that their bases are more or less contiguous (Text-fig. 5) c 10. —. Vertical setae further apart, their bases distinctly: eqpemded ¢ i.e. at fest the diameter of the setal base apart . : Fi 12. to. Lateral and marginal setae slender, plumose only i in their distal third (Text fig. 5); sternal shield with distinct lines ee 1 HE 1) ; lateral processes of the tectum free (Text-fig. 8) 5 : . Macrocheles muscaedomesticae (Scopoli) a Except i in Macrocheles submotus Falconer in which Mgz4 is simple and Mg5 and 6 pilose, * This species is characterised by having nine pairs of setae in the D series, 12 BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH —. Lateral and marginal setae strongly plumose in their distal two-thirds; lateral processes of the tectum partially fused . - II. 11. With three pairs of platelets between ventri- anal and genital shields (PL. 2 fie. Oo Macrocheles cavinatus (C. L.Koch) —. Without platelets between ventri-anal and genital shields ; : - ‘ Macrocheles penicilliger (Berl.) 12. Setae D2, Mg2 and Mgy simple (Text-fig. 36) é 4 Macrocheles submotus Falconer —. Setae D2, Mg2 and Mg4 plumose 9 13% 13. Setae Mr simple, approximately equal in length to Di and extending beyond the bases of D2 by about one-half their length (Text-fig. 42) Macrocheles tayvdus (C. L. Koch) —. Setae Mi simple or plumose and considerably shorter in length than Dr 2 é 14. 14. Dorsal shield less than 950z in length - 5 : 5 2 6 : 4 15. Dorsal shield more than 1100z in length. 9 16. 15. Setae Mgio, approximately equal in length to DS, are ‘about half the length of Mgo (Text-fig. 44); external margin of the lateral processes of the tectum smooth anterior to the base of the median process (Text-fig. 45) ; punctate areas on the sternal shield inconspicuous . ; Macrocheles decolorvatus (C. L. Koch) —. Setae Mgto equal in length to Mgo (Text: me 47); external margin of the lateral processes of the tectum serrate well beyond the base of the median process (Text- fig. 48) ; punctate areas on the sternal shield large, conspicuous Macrocheles matrius Hull 16. Setae Mr plumose and lying in line with setae D1 (Text-fig. 50) ; lateral margins of the dorsal shield coarsely and unevenly serrated (Text-fig. 51) ; ventri-anal shield considerably broader than long and characteristically shaped (PI. 3, fig. 15) Macrocheles plumiventris Hull —. Setae Mr simple and lying in line with D2 (Text-fig. 56) ; lateral margins of the dorsal shield minutely and evenly serrated re fig. 5 ay ventri-anal shield considerably longer than broad : : . Macrocheles superbus Hull Macrocheles muscaedomesticae (Scopoli) Sellnick. Acarus muscae domesticae Scopoli, J. A. (1772). Annus. V. Hist. Nat. :n. 125, 157. Acarus marginatus Hermann, J. F. (1804). Mém. Apt. : 76, figs. Macrocheles muscae domesticae, Sellnick, M. (1940). Géteborg. Vetensk. Samh, Handl. (5) 6B, No. 14: 78, figs. Macrocheles muscaedomesticae, Pereira, C. & de Castro, M. P. (1945). Arg. Inst. Biol. S. Paulo 6: 163, figs. Female. Dorsal shield reticulated and bearing twenty-eight pairs of setae and twenty-two pairs of “ pores ’”’ (Text-fig. 5). Vertical setae plumose in their distal third and lying in close proximity to each other. Setae D5—D7, M1, M3 and Mgq simple. The remainder of the setae on the dorsal shield are pilose in their distal third. The distribution of the setae and pores is shown in the figure. Extra-marginal setae are simple (Text-fig. 6). The sternal shield is characteristically ornamented with punctures and ridges (Pl. 1, fig. 1.) All the sternal setae are simple. The metasternal shields are free and each carries a simple seta. Genital shield, truncate posteriorly, bears a simple pair of setae and is ornamented with punctate lines. The ventri-anal shield (approx. 368 x 379) has a loose network of punctate lines. All the setae on this compound shield are simple. The peritrematal shield is free posteriorly ; being separated from BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH 13 the exopodal shields by a wide expanse of striated cuticle (Text-fig. 6). The peritreme extends beyond coxa I. The gnathosoma is strongly sclerotized and carries four pairs of setae ventrally (Text-fig. 7). The ventral groove is provided with six or seven transverse rows of denticles. Anteriorly the corniculi and salivary stylets are strongly developed. The chaetotaxy of the five free segments of the pedipalp is normal for the free-living Gamasina. The specialized seta on the inner basal angle of the palptarsus is three- pronged. The lateral processes of the tectum are free ; the median process is bifid distally (Text-fig. 8). The chelicerae are massive ; the movable digit being tridentate and the fixed bi- or tridentate (Text-fig. 9). Leg I has the tarsus (187) longer than the tibia (165). Male. This sex and the immature stages are described and figured by Pereira & de Castro (1945). Dimensions. Female: length 980-1014; breadth 570-6714. Male: length 750-goou ; breadth 450—600p. HABITAT AND LOCALITY. This species is commonly found on Musca domestica Linn. and allied species. Pereira & de Castro (1945) state that all instars except the larval feed on the eggs of house flies. M. muscaedomesticae is cosmopolitan in distribution. Macrocheles pisentii (Berl.) Gamasus tavdus var. Pisentii Berlese, A. (1882). Bull. Soc. ent. Ital. 14: 112, fig. Holostaspis Pisentii Berlese, A. (1887). Acari, Myriopoda et Scorpiones, etc., Fasc. 76, N. 1. Female. Dorsal shield, minutely punctured, and bearing twenty-eight pairs of setae. Vertical setae simple and well separated from each other (Text-fig. 10). Setae D8 short and comb-like (Text-fig. 11). The remainder of the setae on the dorsal shield long-and sharply pointed distally. Extra-marginal setae simple. Ventrally, the sternal shield (without distinct ornamentation) has three pairs of simple setae. Its posterior margin is strongly concave. Metasternal shields small ; setae simple. Genital shield with well-developed lateral sclerites, Ventri-anal shield (about 285 x 220) is considerably longer than broad. All the setae on this shield are simple. External posterior rostrals are about-one-third the length of the internals. The ventral groove has five transverse rows of denticles. The characteristic form of the tectum is shown in Text-fig. 12. The fixed digits of the chelicerae are provided with one large, grooved tooth and five or more small teeth (Text-fig. 13). The movable digit is tridentate ; the middle tooth being large and recurved. Leg I has the tarsus (143) longer than the tibia (132). Tarsus II is provided with stout spines. Male. Unknown. Dimensions. Female: length 810-835; breadth 490-506y. HABITAT AND LOCALITY. This species has been collected from Scarabaeus semt- punctatus in Italy. Hull (1918) records it from a nest of Formica rufa at Chopwell, 14 BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH Durham ; from moss at Ninebanks, Northumberland ; and in dead leaves, Cheshire. We have not examined Hull’s specimens. The above description and figures are based on specimens in the Oudemans Collection, Leiden. wot ba I3 Fics. 10-13. Macrocheles pisentii (Berl.), female. Fig. 10, dorsal shield; Fig. 11, seta D8 enlarged. Fig. 12, tectum. Fig. 13, chelicera. BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH 15 Macrocheles rothamstedensis sp. nov. FEMALE. Dorsal shield, with twenty-eight pairs of setae and twenty-two pairs of “pores ”’, is weakly ornamented with punctate lines forming a polygonal network (Text-fig. 14). The vertical setae are long and slightly pilose distally ; their bases are almost contiguous (Text-fig. 15). Setae D4, Lx, Lz, L6, Mgr and Mgro are conspicuously pilose distally, other dorsal setae may show slight pilosity distally. All the extra-marginal setae are simple. 16 Fics. 14-16. Macrocheles vothamstedensis sp. nov., female. Fig. 14, dorsal shield. Fig. 15, setae Dx enlarged. Fig. 16, chelicera. 16 BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH The sternal shield is densely covered with punctures which tend to form a pattern of punctate lines (Pl. 1, fig. 2). All the sternal setae are simple. The smooth meta- sternal setae lie on small narrow shields. The genital shield, with a pair of simple setae, is truncate posteriorly and ornamented with a network of punctate lines. The lateral sclerites are strongly formed. The ventri-anal shield (approx. 250 x Ig8y) is longer than broad and ornamented with a loose network of punctate lines. All the setae on this shield are simple. The metapodal shields are small and weakly sclerotised. The venter of the gnathosoma is minutely punctured. The ventral groove is | provided with five rows of denticles. Each lateral process of the tectum is free. The movable digit of the chelicera is tridentate; the fixed digit is basically bidentate (Text-fig. 16). Leg I (approx. 610m in length) with the tibia (troy) evidently shorter than the tarsus (143). Legs II and III with simple setae ; leg 1V with plumose setae on the femur, genu, tibia and tarsus. Mate. Dorsal shield, strongly attenuated posterior to coxae IV, bears thirty pairs of setae. The chaetotactic pattern differs from that in the female in the addition of two pairs of extra-marginal setae to the shield and the greater distance between the verticals. The ornamentation of the shield is similar to that in the female. The venter is covered by a punctured holoventral shield bearing nineteen simple setae. The tectum is basically the same as in the female. The fixed digit of the chelicera is tridentate and the movable is provided with two or three small teeth. The sperma- tophoral process is about the length of the movable digit. Femur, genu and tibia of leg II and the femur and tibia of leg IV are spurred. Femur II and trochanter IV have small sclerotized ridges. Dimensions. Male: length 590-595; breadth 365-370”. Female; length 710-7404 ; breadth 370-375. HABITAT AND LOCALITY. Seven females and three males from bullock manure, Rothamsted Experimental Station, Harpenden, Herts. Holotype female, 1955.10. 22.43; allotype male, 1955.10.22.44 and paratypes, 1955.10.22.45-52. Macrocheles glaber (Miiller) Holostaspis glabya Miiller, J. (1860). K. K. méhyr. schles. Ges. Briinn : 178, figs. Gamasus steycovarius Kramer, P. (1876). Arch. Naturgesch. 42: 95, fig. Holostaspis badius, Berlese, A. (1889). Acari, Myviopoda etc., fasc. 52, N. 3. Macrocheles marginatus var. littorvalis Halbert, J. N. (1915). Proc. Roy. Ivish Acad. 31, 39 ii: 67, fig. Macrocheles (Coprholaspis) glabey Berlese, A. (1921). Redia 14: 85. \Macrocheles (Monoplites) oudemansii Hull, J. E. (1925). Ann. Mag. nat. Hist. (9) 15: 215. (in part) Macrocheles veterrimus Sellnick, M. (1940). Géteborg. Vetensk. Samh. Handl. (5) 6B : 80, figs. Coprholaspis anglicus Turk, F. A. (1946). Ann. Mag. nat. Hist. (11) 12: 791, figs. syn. nov. 1 Hull proposed this name for Macrocheles marginatus Oudemans, 1901 nec Herman, 1804. This ““ species,”’ however, is a complex of at least two distinct species: the tritonymph being M. glaber and the adult, M. plumiventris. BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH 17 18 Fics. 17-20. Macrocheles glaber (Miiller), female. Fig. 17, dorsal shield. Fig. 18, setae Di enlarged. Fig. 19, tectum. Fig. 20, chelicera. ZOOL, 4, I. 2 18 BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH FEMALE. The dorsal shield is weakly reticulated and punctured, and carries twenty-eight pairs of setae and twenty-two pairs of “‘ pores” (Text-fig. 17). The vertical setae, lying in close proximity to each other, are plumose in their distal half. Setae D8 are finely pilose, and D4 and Lz2 spiculate distally. The remainder of the setae on the shield are smooth and sharply pointed. The extra-marginal setae are simple and curved. The tritosternum is normal. The sternal shield is characteristically ornamented with lines and punctate areas (PI. 1, fig. 3). The transverse anterior, arcuate, trans- verse median and oblique lines are well defined. All the sternal setae are simple. The metasternal shields are small and the setae simple. The genital shield is strongly ornamented and provided with well-developed accessory sclerites. The ventri-anal shield (approx. 264-275 long x 265-295 wide) is ornamented with concentric punctate lines. The median pre-anal setae are situated well outside the connecting line between the anterior and posterior pre-anals. All the setae on the ventri-anal are simple. The metapodal shields are irregular in outline. The peritreme and peri- trematal shield are normal for the genus. The gnathosoma bears four pairs of setae ventrally. The internal posterior rostrals are about four times the length of the externals. The ventral groove has six transverse rows of denticles. The pedipalps are normal. The tectum is produced into three distinct processes ; the median being divided distally (Text-fig. 19). The chelicerae are strongly developed. The fixed digit is bidentate with the proximal tooth large and ridged (Text-fig. 20). The pilus dentilis is relatively short and stout. The movable digit is tridentate. The dorsal seta is comb-like distally. Leg I (585) has plumed setae on the dorsal surface of the femur and genu. The tibia and tarsus are approximately equal in length (about 115). Legs II (510), III (460) and IV (760) have a few plumose setae on the femur and tarsus. Mate. This sex is figured by Berlese (1889). Dimensions. Female: length 850-855; breadth 540-560y. HABITAT AND LocaLity. This is one of the commonest species of mites found on “dor ’’ beetles in Europe, especially on Geotrupes stercorarius Linn. It is also com- monly encountered on Muscid flies. Macrocheles punctoscutatus sp. nov. Femate. Dorsal shield densely covered with minute punctures and bearing twenty-eight pairs of setae and twenty-two pairs of “‘ pores’”’ (Text-fig. 21). Vertical setae coarsely plumose, well separated (Text-fig. 22). The majority of dorsal setae smooth and sharply pointed apically ; setae D8, Mg2, Mg8-ro distinctly pilose. Extra-marginal setae simple. Tritosternum normal, lacinae long and pilose. Sternal shield heavily ornamented, punctate areas conspicuous (PI. 1, fig. 4). All sternal setae simple. Metasternal shields small, metasternal setae simple. Genital shield strongly ornamented, genital setae simple. Ventri-anal shield (320% * 355) broader than long with its anterior margin lying in close proximity to the genital shield. The ornamentation of the BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH 19 shield consists of distinct lines forming a loose network, and numerous small punctures. The pre-anals, para-anals and post-anal seta are simple. The metapodals are small and elongate ; their long axis lying more or less parallel with that of the longer axis of the body. Mg2 L1 22 M1 Fics. 21-22. Macrocheles punctoscutatus sp. nov., female. Fig. 21, dorsal shield. Fig. 22, setae Dr and Mr enlarged. The external posterior rostral setae are about one-half the length of the internals. The ventral groove has five transverse rows of denticles. The dentition of the chelicerae and the structure of the tectum are similar to that in M. glaber. Leg I (605) has the tibia and tarsus of about equal length. Dimensions. Length 880”; breadth 605. HABITAT AND LocALITy. A single female (holotype 1955.10.22.70) from a mole’s nest at Churcham, Gloucestershire (Coll. R. S. George). Macrocheles subbadius (Berl.)! Holostaspis subbadius Berlese, A. (1904). Redia 1: 264. Female. Dorsal shield bears twenty-eight pairs of setae and twenty-two pairs of “ pores’ (Text-fig. 23). The vertical setae are plumose distally. Setae D8 and, 1 This species is figured by Berlese (1889) under Holostaspis marginatus (Herm.) Berl. ‘‘ forma inter- media inter badium et adultum (marginatum) foem "’ in Acari Myriopoda, etc., fasc. 52, N. 6. 20 BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH in the majority of the specimens examined, setae L2 are pilose. The extra-marginal setae are simple. The sternal shield is ornamented with distinct punctures arranged as in PI. 1, fig. 5. The sternal, metasternal and genital setae aresimple. The ventri-anal shield (190-220 Igo) is ornamented with punctate lines ; the nine setae on this shield are simple. The metapodal shields are weakly sclerotized. OE Fic. 23. Macrocheles subbadius (Berl.), female. Dorsal shield. The lateral processes of the tectum are free and the form of the chelicerae is essentially the same as in M. merdarius. Leg I has the tibia (82) shorter than the tarsus (trom). The spines on tarsus II are short and stout. Mate. Unknown. Dimensions. Length 610-6254; breadth 370-400p. BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH 21 HABITAT AND LOCALITY. Berlese (1889) records this species from manure in Italy. The writers have examined specimens from farmyard manure, Evesham, Worcestershire. Macrocheles insignitus Berl. Macrocheles (Coprholaspis) insignitus Berlese, A. (1918). Redia 13: 158. FEMALE. Dorsal shield with twenty-eight pairs of setae and twenty-two pairs of “pores ’’. The surface of the shield is covered with minute punctures which form a polygonal network. All the setae on the shield and the interscutal membrane are simple. The verticals are short spine-like. Sternal shield characteristically ornamented with punctures as in PI. 1, fig. 6. All the sternal setae are simple. The metasternals are small and flank the anterior part of the ornamented genital shield. The metasternal and genital setae are simple. The ventri-anal shield is about as broad as long (156 x 152) and ornamented with punctate lines. The setae on this compound shield are all simple. The metapodals are extremely weakly sclerotised. The gnathosoma is typical for the genus. The lateral processes of the tectum are free and fish-tail like ; the median process is strongly bifurcate distally. The form of the chelicerae is similar to that in M. merdarius. Leg I (approx. 350m in length) has the tibia (55) considerably shorter than the tarsus (72). The male is unknown. Dimensions. Length 445u; breadth 275. HABITAT AND LOCALITY. This species is previously known by a single female from the type locality ‘‘ Longny, Orne in Gallia’’ (Berlese, 1918). The writers have examined a single female of this species from “a hot bed”’, Austrey, Warwickshire (undetermined in the Michael Collection). Macrocheles merdarius (Berl.) Holostaspis merdarius Berlese, A. (1889). Acari, Myriopoda et Scorpiones etc., Fasc. 52, N. 1, fig. Macrocheles merdarius, Sellnick, M. (1940). Gdéteborg. Vetensk. Samh. Handl. (5) 6B : 86, figs. FEMALE. Dorsal shield, ornamented with reticulations, is provided with twenty- eight pairs of simple setae and twenty-two pairs of pores (Text-fig. 24). The vertical setae are sub-spinose. The extra-marginal setae are also simple. The sternal shield is lightly ornamented with punctate lines and the three pairs of sternal setae are simple (PI. 2, fig. 7). The metasternal shields are minute. Both the metasternal and genital setae are simple. The truncated anterior margin of the ventri-anal shield lies in close proximity to the posterior margin of the genital. The ventri-anal (about 150 x 126) is ornamented with four or five transverse lines. All the setae on this shield are simple. The metapodals are small and weakly sclerotized. 22 BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH Ventral groove with five transverse rows of denticles. External posterior rostrals about one-third the length of the internals. The tectum has the lateral processes free (Text-fig. 25). The median process is deeply bifurcate. The dentition of the digits of the chelicerae is shown in Text-fig. 26. oP 56) Fics. 24-26. Macrocheles merdarius (Berl.), female. Fig. 24, dorsal shield. Fig. 25. tectum. Fig. 26, chelicera. Mate. This sex is described and figured by Berlese (1889). Dimensions. Female: length 445-490”; breadth 225-280u. Male: length 380 ; breadth not given by Berlese (1880). HABITAT AND LOCALITY. This is one of the commonest species of Macrochelids occurring in dung and compost. It has been recorded from a number of localities in a BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH 23 Europe (Franz, 1954). In Britain it is recorded from bullock dung at Rothamsted Experimental Station, Harpenden, Herts. (Hyatt, 1956). Macrocheles montanus (Willmann) Nothrholaspis montana Willmann, C. (1951). Bonner Zool. Beitr. 2: 158, figs. FEMALE. Dorsal shield with twenty-nine pairs of setae and twenty-two pairs of “pores ’’ (Text-fig. 27). The surface of the shield is conspicuously punctated and Fic. 27. Macrocheles montanus (Willmann), female. Dorsal shield. 24 BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH reticulated. Dorsal series (D) comprising nine pairs of setae. Vertical setae strongly plumose with their bases well separated. Setae Mr, D5—D8, M3 and M4 and the additional setae of the D series are simple. The remainder of the dorsal setae are plumose. Extra-marginal setae are simple. The sternal shield is covered with punctures which are larger in the posterior third of the shield (Pl. 2, fig. 8). The first pair of sternal setae are plumose in their distal third ; the second and third pairs are smooth or slightly plumose distally. The form of the metasternal and genital shield is shown in the figure. Ventri-anal shield (357 x 28 Fic. 28. Macrocheles montanus (Willmann), female. Chelicera. 3401) is ornamented with a network of punctate lines. The median pre-anal seta lies almost on the line connecting the anterior and posterior pre-anals. The pre-anal and para-anal setae are simple, the post-anal seta is strongly plumose distally. There are three pairs of platelets between the genital and ventri-anal shields. The metapodal shields are weakly sclerotized. Venter of the gnathosoma normal for the genus. Tectum essentially the same as in M. submotus (Text-fig. 37). Both digits of the chelicerae tridentate (Text-fig. 28). Leg I with the tarsus (175) longer than the tibia (154,). The male is unknown. Dimensions. Length 1,050-1,1404 ; breadth 660y. HABITAT AND LocaLiTy. This species is previosuly known from Austria only BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH 25 (Willmann, 1951 and Franz, 1954). We have examined a single female collected under “ old wet oak bark ’’ in Marley Wood, Wytham, Berkshire (Coll. E. W. Fager). M.montanus is the only British species in the genus which has more than the normal compliment of setae on the dorsal shield. In Willmann’s figure of the type specimen there are thirty-one pairs of setae on the shield; additional plumose setae being present near setae L7 and Lo as well as the simple pair in the D series. In other characters the British specimen agrees with the type. Macrocheles carinatus (C. L. Koch). Gamasus cavinatus Koch, C. L. 1839. Deutsch. Crust. Myr. Avach. fasc. 24, t. 16. Macrocheles hypochthonius Oudemans, A. C. 1913. Ent. Ber. Amst. 4: 6. Macrocheles hypochthonius Oudemans, A. C. 1914. Arch. Naturgesch. 79a, Hft. 8: 175, figs. Nothrholaspis hulli Falconer, W. 1923. Naturalist Lond. : 153, figs. syn. nov. Nothrholaspis carinata Sellnick, M. 1931. SB. Akad. Wiss, Wien 140: 766, figs. FEMALE. The dorsal shield is strongly reticulated and punctured, and its lateral margins serrated (Text-fig. 29). The vertical setae (D1) which stand in close proximity to each other, are long and strongly plumose in their distal half (Text-fig. 30). Setae Mr are short and may be smooth or plumose. Setae M3, M4, D6, D7, and D8 are thin and simple or slightly serrated distally. The remainder of the setae on the dorsal shield are strong and plumose. Setae D3 lie in advance of M2. The anterior extra-marginal setae are simple and the posterior plumose. The sternal shield is ornamented with a network of ridges and punctures ; the latter are especially large in the posterior third of the shield (Pl. 2, fig. 9). The sternal and metasternal setae are simple. The posterior margin of the genital shield is strongly convex, the genital setae are simple. The ventri-anal shield is sub-circular in outline and is strongly ornamented. All the setae on the compound shield are simple. The median pre-anals lie considerably closer to the anterior than the posterior pre-anals and are situated almost on the connecting line between these setae. The interscutal membrane between the genital and ventri-anal shield bears six platelets. The metapodal shields are small. The peritreme and peritrematal shield are normal for the genus. The interscutal membrane is conspicuously corrugated. . The external posterior rostral setae are about one-third the length of the internals. The ventral groove carries five rows of denticles. The tectum (Text-fig. 31) has the lateral lobes partially fused; the lateral and median lobes are divided distally. The chelicerae are strongly developed with the fixed digit tri-dentate (Text-fig. 32). The movable digit may be bi- or tri-dentate. The dorsal seta is serrated on one side. Leg I, approximately 935 in length, has the tibia (120) considerably shorter than the tarsus (165). Legs II, III, IV measure approximately 660y, 6051, and 1,100p respectively. Mate. The chaetotaxy of the dorsal shield is basically the same as in the female. The sterniti-genital shield is provided with a polygonal network of ridges and strong punctures. The ventri-anal shield is also reticulated and punctured and measures about 298m in length and 310m in breadth. All the setae on this shield are simple. 26 BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH The tectum is similar to that in the female. The spermatophoral process on the movable digit of the chelicera is short, being about one-half the length of the digit. Femur, genu and tibia of leg II only are spurred. Dimensions. Male: 880-gooy in length ; 500-540 in breadth. Female: 1,034— 1,078 in length ; 570—-640y in breadth. HABITAT AND LocaLity. In humus and moss from a number of localities in the British Isles. This species has also been recorded from Austria, Germany and the Netherlands. ty DI elle i3) Fics. 29-32. Macrocheles cavinatus (C. L. Koch), female. Fig. 29, postero-lateral margin of the dorsal shield. Fig. 30, anterior region of the dorsal shield. Fig. 31, tectum. Fig. 32, chelicera. BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH 27 Macrocheles penicilliger Berl. Holostaspis penicilligey Berlese, A. (1904). Redia 1: 264. Holostaspis penicilligey Berlese, A. (1918). Redia 13: 146, 162. Macrocheles penicilliger, Sellnick, M. (1940). Gdéteborg. Vetensk. Samh. Handl. (5) 68 : 82, figs. FEMALE. The dorsal shield is strongly reticulated and punctured, and has its lateral margins serrated (Text-fig. 33). Vertical setae, approximately 55 in length, are plumose in their distal two-thirds. Their bases are almost contiguous. Setae Mr, M3, M4, D6—D8 are simple ; remainder of dorsalsetae plumose. Setae D8 are situated considerably anterior to the line connecting setae Mgio. Extra-marginal setae are plumose distally. Fics. 33-35. Macrocheles penicilliger (Berl.), female. F ig. 33, dorsal shield. Fig. 34, tectum. Fig. 35, chelicera. 28 BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH The sternal shield is heavily ornamented with ridges and punctures (PI. 2, fig. 10) The first pair of sternal setae are strongly plumose and the second and third pair slightly plumose distally. The metasternal setae are simple as are the pair of setae situated on the punctured genital shield. On the ventri-anal shield (approx. 310 x 330), the pre-anal and para-anal setae are simple and the post-anal seta plumose. The median pre-anal lies outside the line connecting the anterior and posterior pre-anal. There are no platelets between the ventri-anal and genital shields. The metapodals are strongly sclerotized and elongate. The external posterior rostrals on the venter of the gnathosoma are about one- third the length of the internals. Ventral groove has five transverse rows of denticles. The lateral processes of the tectum are partially fused ; distally these processes may or may not be divided (Text-fig. 34). The median process is deeply bifurcate. Both digits of the chelicerae are bidentate (Text-fig. 35). The pilus dentilis is short and thick at its base. The ventral seta is long and strongly pilose. Leg I with tibia (approx. 99/2) considerably shorter than the tarsus (approx. 130/). Mate. Unknown. Dimensions. Female: length goo—930u ; breadth 525-550y. HABITAT AND LOCALITY. This species has been found on Tvox scaber (Linn.) in the nests of owls at Wytham, Oxford (Coll. C. E. Elton) and at Woodford, Essex (Coll. A. Hooper). Further records are from decaying leaves, Waterworks Valley, Jersey (Coll. G. Owen Evans), Italy (Berlese, 1904), Iceland (Sellnick, 1940) and Austria (Franz, 1954). Macrocheles submotus Falconer, Macrocheles cognatus Falconer, W. (1923). Naturalist, Lond. : 152, fig. (nom. praeocc.). Macrocheles submotus Falconer, W. (1924). Naturalist, Lond. : 363 (nom. nov. pro. M. cognatus). Macrocheles (Nothrholaspis) occidentalis Hull, J. E. (1925). Ann. Mag. nat. Hist. (9) 15: 213, fig. syn. noy, ? Macrocheles (Nothrholaspis) gloriosus Hull, J. E. (1925). Ann. Mag. nat. Hist. (9) 15: 214, fig. syn. nov. FEMALE. Dorsal shield, strongly reticulated and punctured, is provided with twenty-eight pairs of setae and twenty-two pairs of pores. Its lateral margins are entire. Setae D1 plumose and situated about one diameter of their bases apart (Text-fig. 36). Setae Mr, M4, D2, D5—D8, L1!, Mg2 and Mg4 are simple, and Mg5 and Mgé6 pilose. The remainder of the setae on the dorsal shield are distinctly plumose. The extra-marginal setae are simple or slightly pilose. The tritosternum is normal with strongly pilose lacinae. The sternal shield, ornamented with strong ridges and large punctures, bears three pairs of simple, spine-like setae (Pl. 2, fig. 11). The metasternal and genital setae are also simple. Ventri-anal shield (approx. 450 x 430m) is triangular in outline and ornamented with ridges and punctures. The pre-anal and para-anal setae are simple. The median pre-anal lies outside the line connecting the anterior and posterior pre-anals. Three pairs of platelets lie between the ventri-anal and genital shields. The interscutal mem- brane is closely striated. The peritreme and peritrematal shields are normal. 1 This seta should be simple in Text-fig. 36. BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH Fics. 36-39. Macvocheles submotus Falconer. Fig. 36, dorsal shield of female. Fig. 37, tectum of female. Fig. 38, chelicera of female. Fig. 39, chelicera of male. 29 30 BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH The external posterior rostrals on the venter of the gnathosoma are about one-half the length of the internals. The ventral groove carries five transverse rows of denticles The form of the tectum is shown in Text-fig. 37. Both digits of the chelicerae are massive (Text-fig. 38). The fixed digit has two strong and two weaker teeth; the movable digit is basically bi-dentate. The dorsal seta is comb-like. Leg I (approx. 1,100m in length) has the tibia (205) shorter than the tarsus (220). Legs II-IV measure respectively about 1,150, 1,078 and 1,700. Mate. The chaetotaxy and ornamentation of the dorsal shield is essentially the same as in the female. Ventrally, the truncated sterniti-genital shield is coarsely punctured. The five pairs of setae on this compound shield are simple. The ventri- anal shield (350 x 340) is reticulated and punctured ; there are no platelets between the ventri-anal and genital shield. Both digits of the chelicerae are dentate as in Text-fig. 39. The spermatophoral process is considerably shorter than the length of the digit. The femur, genu and tibia of leg II only are spurred. Dimensions. Male: length 1,050-1,100”; breadth 620-630”. Female: 1,390- 1,450; breadth 780-82o0p. HABITAT AND LOCALITY. This is one of the commonest, and most widely distributed Macrochelid found in litter and humus under deciduous and coniferous trees in Britain. The above description of swbmotus is based on specimens compared with the type in the Falconer Collection at the Liverpool Museum. This species has also appeared under the name of Macrocheles tridentinus (Can.) in British and possibly other European faunal lists. The original description of trzdentinus was based on the male only and both description and drawings are insufficient for the certain identity of the species. Canestrini gives the length of the male of tridentinus as 860. Macrocheles tardus (C. L. Koch) Gamasus tavdus Koch, C. L. 1841. Deutsch. Crust. Myr. Arach. fasc. 39, t. 14. Nothrholaspis tardus, Sellnick, M. 1931. S.B. Akad. Wiss. Wien. 140: 765, figs. FEMALE. Dorsal shield, strongly reticulated and punctured, is provided with twenty-eight pairs of setae and twenty-two pairs of pores. The vertical setae (Dr) are plumose and well separated, setae Mr are long, simple and extend beyond the bases of D2 by about one-half their length (Text-fig. 40). Setae M3, M4, D6 and D8 are simple, D7 are finely pectinated whilst the remainder of the setae of the dorsal shield are strongly plumose. The marginal setae (Mg) are situated a short distance from the lateral margin of the shield (Text-fig. 41). The anterior extra-marginal setae are pectinate and the posterior extra-marginals plumose. The tritosternum is normal with strongly pilose lacinae. The sternal shield is ornamented with a polygonal network of ridges and numerous punctures (Pl. 2, fig. 12). All the sternal setae are simple. The metasternal shields are small and the setae plumose. The strongly punctured genital shield is convex posteriorly and bears a pair of simple setae. The ventri-anal shield, about as broad as long (330 x 300/) is BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH 31 reticulated and punctured. The anterior and posterior pre-anals are simple but the median pre-anal may be simple or pectinate. The para-anals are long and simple and the post-anal seta short and plumose. The region between the genital and ventri-anal shields is usually provided with three pairs of platelets. This number may be reduced in some specimens. The metapodal shields are small. The peritreme and peritrematal shield are normal. The interscutal membrane is coarsely striated, the striae being provided with triangular processes at intervals along their length. Ventrally, the gnathosoma has the normal four pairs of setae of which the external posterior rostrals are about one-half the length of the internals. The ventral groove y, Fics. 40-43. Macrocheles tavdus (C. L. Koch), female. Fig. 40, anterior region of the dorsal shield. Fig. 41, postero-lateral margin of dorsal shield. Fig. 42, tectum. Fig. 43, chelicera. 32 BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH has five rows of denticles. The lateral processes of the tectum are partially fused (Text-fig. 42) ; the distal end of these processes being bi- or trifurcate. The stout median process is divided ; each arm being entire or divided distally. The fixed digit of the chelicera is provided with two strong teeth and between them two or three weaker teeth (Text-fig. 43). The pilus dentilis is long and stout. The movable digit has two recurved teeth only. The dorsal seta is large and comb-like. Leg I (approximately 1,020 in length) bears plumose setae on the femur, genu and tibia. The tibia (165-175) is shorter than the tarsus (200-215). Leg II (880) has plumose setae on the trochanter and tarsus as have leg III (820) and leg IV 1,485). Mate. The chaetotaxy and ornamentation of the dorsal shield is essentially the same as in the female. The sterniti-genital shield, truncated posteriorly, is heavily reticulated and punctured. The five pairs of setae on the compound shield are simple. The ventri-anal shield (330 x 350j) is also reticulated and punctured. The pre-anal and para-anal setae are usually simple, but may be pectinate. The post- anal seta is strongly pilose. The chelicerae are basically the same as in M. cognatus, differing only in the position of the large proximal tooth on the fixed digit. The spermatophoral process is considerably shorter than the length of the movable digit. The femur, genu and tibia of leg II are the only segments which are spurred. Dimensions. Male: 1,030-1,100~% in length; 627—7oou in breadth. Female I,215-1,290y in length ; 730-800 in breadth. : HABITAT AND Locality. In vegetable debris near Rydal Water, Westmorland and in decaying leaves, Waterworks Valley, Jersey. This species has also been recorded from a number of other localities in Europe (Franz, 1954). Macrocheles decoloratus (C. L. Koch) Gamasus decolovatus Koch, C. L. (1893). Deutsch. Crust. Myr. Arach. fasc. 25, t. 14. Macrocheles decolovatus, Oudemans, A. C. (1913). Ent. Ber. Amst. 4: 5. Macrocheles decolovatus, Oudemans, A. C. (1914). Arch. Naturgesch. 79a Hft. 8 : 173. FEMALE. Dorsal shield, weakly reticulated but strongly punctured, bears twenty- eight pairs of setae and twenty-two pairs of pores (Text-fig. 44). The vertical setae (Dr) are well separated and strongly plumose. The remainder of the setae on the dorsal shield are also plumose. A characteristic feature of the chaetotaxy is the relative length of setae Mgg and 10. The latter is approximately equal in length to D8, but only about one-half the length of Mgg. The extra-marginal setae are plumose. The tritosternum is normal. The sternal shield (195 long x 220m wide) is orna- mented with a faint reticulate pattern and large punctures (PI. 3, fig. 13). The punc- tate areas are usually conspicuous. The three pairs of sternal setae and the metaster- nals are long and simple or slightly pectinate. The genital shield is strongly punctured and has a network of punctate lines in its anterior half. The genital setae usually are simple. The ventri-anal shield (286-298 long x 250-286 wide) is provided with a network of punctate lines. The lateral regions of the shield are densely punctured. The pre-anal and para-anal setae are long and simple. The post-anal seta is shorter and plumose. There are no platelets between the genital and ventri-anal shield. The BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH 33 metapodals are small and irregular in outline. The peritreme and peritrematal shields are normal. Ventrally, the gnathosoma carries the normal four pairs of setae. The external posterior rostrals are about one-third the length of the internals. The capitular Fic. 44. Macrocheles decolovatus (C. L. Koch), female. Dorsal shield. groove is provided with five transverse rows of denticles. The tectum (Text-fig. 45) has the three anterior processes separate. The movable digit of the chelicera is tridentate and the fixed bi- or tridentate (Text-fig. 46). The dorsal seta is spatulate. Leg I (approx. 670m) with plumose setae on femur, genu and tibia. Tarsus I (132-140) longer than tibia I (x21). Leg II (680u). Leg III (638) and Leg IV (990) with some plumose setae on trochanter and tarsus. Mate. Unknown. ZOOL. 4, I. 34 BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH Dimensions. Length 850-880 ; breadth 510-520. HABITAT AND LOCALITY. In cow dung, nr. Canterbury, Kent (Coll. E. Warren, 1942). This species has also been recorded from Austria, Germany and the Nether- lands. 45 Fics. 45—46. Macyocheles decolovatus (C. L. Koch), female. Fig. 45, tectum. Fig. 46, chelicera. Macrocheles matrius (Hull). Nothrholaspis matrius Hull, J. E. (1925). Ann. Mag. nat. Hist. (9) 15: 212. Macrocheles subbadius var. vobustulus, Sellnick, M. (1940). Gdteborg. Vetensk. Samh. Handl. (5) 6B : 86, figs. Macrocheles cavinatus, Hughes, A. M. (1948). Mites associated with stored food products. H.M.S.O. : 126, figs. FEMALE. Superficially, the chaetotaxy of the dorsal shield in this species resembles that in M. decoloratus. The chief difference between the species lies in the relative lengths of setae Mgg and Mgto which in the present species are approximately equal in length (Text-fig. 47). The extra-marginal setae are slightly plumose. The sternal shield (187 long x 220s wide) has well developed lines and a few punctures (Pl. 3, fig. 14). The punctate areas are considerably larger and more strongly developed than in M. decoloratus. The sternal and metasternal setae are simple or slightly pectinate. The genital shield is ornamented with punctate lines and large punctures. The genital setae are plumose distally. The ventri-anal shield BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH 35 (3404 < 3754 wide) is characteristically ornamented. The pre-anal and para-anal setae are simple. The post-anal seta is short and plumose distally. Metapodals are elongate and weakly sclerotized. The gnathosoma is essentially the same as in decoloratus except for the structure of Mg10 4 Fic. 47. Macrocheles matrius Hull, female. Dorsal shield. the tectum; in matrius the external margin of the lateral processes are strongly : serrulate (Text-fig. 48). y Mate. This sex is adequately described and figured by Hughes (1948). 3 Dimensions. Male: length 680-720u ; breadth not stated by Hughes (1948). Female: length 890-910”; breadth 540-565. HABITAT AND LOCALITY. Hull (1925) states that the species is ‘‘ not uncommon in north of England. Abundant in poultry manure, West Allendale”. We have 36 BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH examined specimens from the nest of Riparia riparia at Tirley, Gloucestershire (Coll. RS. George) and from poultry manure at Houghton, Huntingdonshire (Coll. C. Horton Smith). Hughes (1948) records it under the name M. carinatus from floor debris and sievings of grain. 48 Fic. 48. Macrocheles matrius Hull, female. Tectum. M. matrius is undoubtedly conspecific with Macrocheles subbadius var. robustulus (Berl.) Sellnick (1940). If the species described and figured by Sellnick is the same as vobustulus, there is some doubt concerning this, then matvius must be considered a synonym. Macrocheles plumiventris Hull Holostaspis marginatus Berlese, A. (1889). Acari, Myriopoda et Scorpiones, etc., fasc. 52, No. 4 and 5, figs. ? Macrocheles gladiatoy Hull, J. E. (1918). Trans. Nat. Soc. Northumb., N.s. 5: 71, figs. ? Macrocheles plumipes Hull, J. E. (1918). tom. cit. : 72, fig. Macrocheles (Monoplites) oudemansii Hull, J. E. (1925). Ann. Mag. nat, Hist. (9), 15: 215 in part). ae plumiventris Hull, J. E. (1925). Ann. Mag. nat. Hist. (9), 15: 216, figs. Nothrholaspis fimicola Sellnick, M. (1931). S.B. Akad. Wiss. Wien 140: 765, fig. syn. nov. FEMALE. The dorsal shield is strongly reticulated, especially in its posterior two-thirds. The lines of the mesh-work are distinctly crenulated (Text-fig. 49). The vertical setae (D1) which are well separated, and setae D2—D4, M2, series L and series Mg. are strongly plumose in their distal half (Text-fig. 50). Setae Mr are short and relatively weakly plumose ; setae D6—D8 and Mg2-3 are considerably finer and less plumose than the L and the remainder of the Mg. series. The marginal setae are situated on the serrated lateral margin of the dorsal shield (Text-fig. 51). The extra-marginal setae are plumose. The tritosternum is normal. All the ventral shields are strongly reticulated and BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH 37 S Bs 54 49 52 dorsal shield. Fig. 50, anterior region of the dorsal shield. Fig. 51, lateral margin of the dorsal shield. Fig. 52, tectum. Fig. 53, chelicera. Fig. 54, dorsal seta on chelicera | Fics. 49-54. Macrocheles plumiventris Hull, female. Fig. 49, ornamentation of the 38 BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH punctured (Pl. 3, fig. 15). The sternal shield bears three pairs of sternal setae of which ht is strongly plumose and h2—3 smooth and blunt. The metasternal shields are free and the setae plumose. The genital shield is provided with accessory sclerites and the genital setae are plumose. The ventri-anal shield is considerably broader than long (370-460 x 490-610) and characteristically shaped. The three pairs of pre- anal setae are plumose, the para-anal setae fine and slightly pilose, and the post-anal seta short and strongly plumose. The metapodal shields are well developed. The peritreme and peritrematal shield are normal for the genus, Ventrally, the gnathosoma bears four pairs of setae. The external posterior rostrals are more than one-half the length of the internal posterior rostrals. The ventral groove is provided with seven rows of denticles. The pedipalps are normal. The tectum has its lateral processes partially fused. The apex of these processes may be entire or bifurcated (Text-fig. 52). The median process is strongly bifid distally. The fixed digit of the chelicera has five teeth of which the larger are ridged (Text-fig. 53). The movable digit is provided with three teeth, the proximal being the smallest. The dorsal seta has a denticulate margin (Text-fig. 54). The ventral setae are long and strongly pilose. Leg I, about 1,200u in length, has plumose setae on the femur, genu and tibia. Trochanter I bears a short conical spur internally. Tibia I measures 220-242 and Tarsus I 210-2204. Leg II is stout, approximately 1,1004 in length and richly provided with plumose setae as are leg III (approx. 1,030~) and Leg IV (approx. 1,650u.). The pulvilli and claws of Legs II-III are well developed. Mate. The ornamentation and chaetotaxy of the dorsal shield is basically the same as in the female. The venter is covered by a reticulated holoventral shield. The fixed digit of the chelicera is provided with four teeth in its distal half. The movable digit, in the specimen under study, has a single tooth. Berlese (1889) shows four teeth on this digit! The spermatophoral process is longer than the movable digit and is pointed distally. The trochanter, femur, genu, tibia and tarsus of Leg IT are provided with spurs, also genu III, and trochanter and femur IV. Dimensions. Male: 913 in length and 605y in breadth (a single specimen in the Michael Collection). Female: 1,150-1,480y in length and 750-935 in breadth. HABITAT AND LocaLity. In manure and compost heaps. Common and widely distributed in Europe. Sellnick (1931) drew attention to the fact that Holostaspis marginatus Berl. (1889) was not Acarus marginatus Hermann (1804) and proposed the name Nothrholaspis fimicola for Berlese’s species. Hull (1925) had, however, already described this species under Macrocheles plumiventris. It is also possible that Hull’s Macrocheles gladiator and plumipes are referable to this species, but the original descriptions and figures are wholly inadequate for their certain identity. Macrocheles superbus Hull Macrocheles superbus Hull, J. E. (1918). Tvans. Nat. Hist. Soc. Northumb., N.s. 5: 71, figs. FEMALE. Dorsal shield minutely punctured and with a faint reticulated pattern in its posterior half. The lateral margin of the shield is evenly serrated (Text-fig. 55). BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH 39 Fics. 55-62. Macrocheles superbus Hull. Fig. 55, lateral margin of dorsal shield. Fig. 56, dorsal shield of female. Fig. 57, tectum of female. Fig. 58, chelicera of female. Fig. 59, distal end of tectum of male. Fig. 60, chelicera of male. Fig. 61, leg II of Male. Fig. 62, leg IV of male. 40 BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH The vertical setae (Dr) are stout, plumose and widely separated (Text-fig. 56). Setae D4—D8, M3 and M4 are simple and minutely spiculate distally. The remainder of the dorsal setae are stout and plumose. The extra-marginal setae are plumose distally. The tritosternum is normal with the lacinae strongly pilose. The sternal shield is ornamented with a network of ridges and punctures (PI. 3, fig. 16). The punctate areas are distinct. The first pair of sternal setae (hi) are strongly plumose whereas h2 and hg are simple and blunt apically. The metasternal setae are similar in form to setae h2 and h3. The genital shield, strongly punctured, is provided with accessory sclerites and a pair of rod-like setae which are spiculate dorsally. The ventri-anal shield (approximately 580 x 460) is strongly reticulated and punctured. The anterior pre-anal setae are simple and the median and posterior setae plumose. The para-anals are simple and the post-anal plumose. A pair of well-sclerotized platelets lie between the genital and ventri-anal shields. The metapodal shields are well developed. The peritreme and peritrematal shield are normal. The four pairs of ventral setae on the gnathosoma are simple with the external posterior rostrals about one-eighth the length of the internals. The former lie well in advance of the latter. The ventral groove is provided with five rows of denticles. The tectum consists of a single process, bifurcate distally (Text-fig. 57). The chelicerae are strongly chelate-dentate with the fixed digit bearing two well-developed teeth and a strong pilus dentilis (Text-fig. 58). The movable digit is also bidentate. Leg I (approx. 480j) with the tibia (230-240) considerably shorter than the tarsus (297-320). Mate. The chaetotaxy and ornamentation of the dorsal shield is essentially the same as in the female except for a characteristic densely punctured area surrounding the base of setae L6. In the majority of the males examined the dorsal shield was strongly attentuated in the posterior third. The sterniti-genital shield is poorly ornamented with a network of ridges and punctures and its posterior margin is slightly concave. The five pairs of setae on the shield are all plumose distally. The ventri-anal shield (approximately 560 x 350) is provided with a polygonal network of ridges and numerous punctations. The pre-anal setae and the post-anal seta are plumose in their distal half. The para-anals are simple. The interscutal membrane between the sterniti-genital and ventri-anal shield is without sclerotized platelets. The metapodal shields are well developed. The structure of the venter of the gnathosoma is basically the same as in the female. The tectum may be bifurcate or more extensively divided distally (Text-fig. 59). The fixed digit of the chelicera is tridentate and the movable bidentate. The spermato- phoral process is about one-third the length of the movable digit (Text-fig. 60). Femur, genu and tibia of leg II (Text-fig. 61) and the trochanter and femur of leg IV (Text-fig. 62) are spurred. Dimensions. Male: 1,420-1,520u in length, 850-goou in breadth. Female: 1450-1650y in length, 790—gooy in breadth. HABITAT AND DISTRIBUTION. Common in wrack and tidal debris above high- water mark on the seashore. Also recorded from salt marshes (Falconer, 1923). Widely distributed in Britain, a - oe BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH 41 SPECIES INCERTAE SEDIS Macrocheles (Nothrholaspis) pannosus Hull Hull, J. E. (1925). Ann. Mag. nat. Hist. (9), 15: 211, fig. It is possible that this species is a small form of M. plumiventris. Dimensions. Female: gooy in length. HABITAT AND LOCALITY. ‘‘ In manure, West Allendale.”’ Macrocheles (Nothrholaspis) nemoralis Hull Hull, J. E. (1925). Ann. Mag. nat. Hist. (9), 15: 213, fig. This species is possibly a synonym of M. penicilliger. Dimensions. Female: 810 in length, 500m in breadth. Hasirat AND Locatity. ‘‘ West Allendale ; habitat unknown.” Macrocheles (Nothrholaspis) parmulatus Hull Hull, J. E. (1925). Ann. Mag. nat. Hist. (9), 15: 214, fig. Dimensions. Female: 1,200y in length. HABITAT AND LOCALITY. ‘‘ West Allendale ; habitat unknown.” Macrocheles (Monoplites) palustris Hull Hull, J. E. (1925). Ann. Mag. nat. Hist. (9), 15: 216, fig. Dimensions. Male: 1,100”; female, 1,200. HABITAT AND LOCALITY. ‘‘ In sphagnum on the moors, West Allendale.” Macrocheles (Monoplites) tardior Hull Hull, J. E. (1925). Ann. Mag. nat. Hist. (9), 15: 216, fig. Dimensions. Female: 1,000y in length. Hasirat AND Locatity. ‘‘ Oxfordshire (R. S. Bagnall). Habitat not stated.” Genus GEHOLASPIS Berl. s. lat. Geholaspis Berlese, A. (1918). Redia, 13: 145. This genus has been revised by Valle (1953) and may be defined as follows : Dorsal shield with twenty-eight pairs of setae and twenty-two pairs of ‘‘ pores”. Sternal shield with three pairs of setae ; metasternal shields free or fused with the endopodal shields. Ventri-anal shield in the female with five pairs of pre-anal setae. Chelicerae normal length with a few teeth or conspicuously elongate and multi- 42 BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH dentate Tectum with median process only. Other characters as in Macrocheles Latr. Type: Gamasus longispinosus Kramer, 1876. Valle (loc. cit.) subdivided the genus into three subgenera of which the following two are represented in the British fauna : 1, Seta Ml long and simple, and extending beyond the bases of D2 by more than half their length (fig. 63). Ratio of the length of the dorsal shield to the length of the movable digit of the chelicera varies between 6°8 and 10°4 ; movable digit with less than 5 teeth . 5 > 2 Geholaspis s. str. Seta MI short, simple or pinniocen and. noe or scarcely: extending beyond the bases of seta D2 (fig. 66). Ratio of the length of the dorsal shield to the length of the movable digit of the chelicera varies between 3'7 and 4°5; movable digit with more than five teeth : a 5 ° - 2 < ; : - Longicheles Valle. The subgenus Geholaspis is represented in Britain by the type species only, and Longicheles by two species, namely, G. (L.) mandibularis (Berl.) and G. (L.) longulus (Berl.). Subgenus GEHOLASPIS Berl. s. str. Geholaspis (Geholaspis) longispinosus (Kramer). Gamasus longispinosus Kramer, P. (1876). Arch. Naturgesch. 42 : too, fig. Holostaspis longispinosus, Berlese, A. (1887). Acari, Myriapoda et Scorpiones, etc., fasc. 44, No. 1, figs. ? Macrocheles castaneus Hull, J. E. (1925). Ann. Mag. nat. Hist. (9), 15: 212. Femae. The dorsal shield is minutely punctured all over and reticulated in its posterior half. It carries twenty-eight pairs of setae and twenty-two pairs of “* pores ”’ distributed as in Text-fig. 63. Setae Di are strongly plumose ; D2, D5—D8, M1, M5 and M6, and Li are simple. The remainder of the dorsal setae are pilose. The extra- marginal setae are similar in structure to the marginal series. The tritosternum is normal with the lacinae pilose. The sternal shield is ornamented with a network of ridges and punctures (Pl. 4, fig. 17). The three pairs of sternal setae and the metasternals are simple. The genital shield is broad and strongly sculptured. The genital setae are simple and the accessory sclerites well-developed. The large ventri-anal shield (440-440 in length and 465-510 in breadth) is broader than long and bears five pairs of pre-anal setae in addition to the three setae normally associated with the anus. The relative lengths of these setae are shown in Text-fig. 4. The surface of the shield is evenly reticulated. The metapodal shields are small and weakly sclerotized. The peritrematal shield is free in its posterior half. Ventrally, the gnathosoma bears the normal four pairs of setae. The external posterior rostrals lie well in advance of the internals. The ventral groove is provided with six transverse rows of denticles. The corniculi are long and slender. The tectum comprises a median process arising from a denticulate base (Text-fig. 64). The process is provided with shoulder-like projections of varying size and shape about a third of the distance from its base ; distally it is divided into two or three pointed branches. a" cers See ae 4 le" BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH 43 The chelicerae are strongly developed and dentate; the dentition of the digits is shown in Text-fig. 65. The ventral setae, omitted from the figure, are well developed. Tibia I (approx. 84) is considerably shorter than tarsus I (approx. 133). Mate. This sex does not appear to have been described. Dimensions. Length, 870-9704 ; breadth 590-710. Fics. 63-65. Geholaspis (Geholaspis) longispinosus (Kramer), female. Fig. 63, dorsal shield. Fig. 64, tectum. Fig. 65, chelicera. HABITAT AND Locality. This species is widely distributed in Europe (Valle, 1953) and relatively common in moss, litter and humus. It has been recorded from North Wales by Hull (1918) and from Ireland by Halbert (1915). The writers have examined specimens from a number of localities in England, Wales and Scotland. The figures of the dorsal and ventri-anal shields of Geholaspis longispinosus (Kramer) given by Valle (1953, figs. I and V (2)) do not appear to refer to this species 44 BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH but to Geholaspis forolivensis Lombardini, 1943, which he considers to be a subspecies of longispinosus. In forolivensis the lateral and marginal series of dorsal setae are strongly plumose as in mandibularis, and the shape of the ventri-anal shield and the relative lengths of the pre-anal setae do not conform with those in longispinosus. Subgenus LONGICHELES Valle The two British species of this subgenus may be separated as follows : 1. Setae Mr simple; a simple seta situated between plumose setae L3 and L4; dorsal shield less than 650. in length a a : : G.(L.) longulus (Berl.) —. Setae Mr strongly plumose ; without simple seta between plumose setae L3 and L4 (Text-fig. 66) ; dorsal shield about 750y in length ° . G.(L.) mandibularis (Berl.) Geholaspis (Longicheles) longulus (Berl.) Holostaspis longulus Berlese, A. (1887). Acavi, Myriapoda et Scorpiones, etc., fasc. 43, no. 9, oe (Longicheles) longulus, Valle, A. (1953). Redia 38 : 351, figs. The key characters given above are based on the description and figures of this species by Valle (1953). The writers have not examined this species, which is recorded by Halbert (1915) from a number of localities in Ireland and by Hull (1918) from Ninebanks, Northumberland. Geholaspis (Longicheles) mandibularis (Berl.) Holostaspis mandibularis Berlese, A. (1904). Redia, 1 : 263. ? Macrocheles minimus Hull, J. E. (1918). Trans. Nat. Hist. Soc. Northumb. n.s. 5 : 73, fig. Geholaspis (Longicheles) mandibularis, Valle, A. (1953). Redia 38 : 344, figs. FEMALE The dorsal shield is densely covered with minute tubercles and bears twenty-eight pairs of setae, of which twenty-two pairs are strongly plumose. The distribution of setae in the posterior half of the shield shows some variation in the material examined. The normal chaetotactic pattern is shown in Text-fig. 66 and a variant in Text-fig. 67. The lateral margins of the shield are conspicuously ser- rated. The vertical setae lie in close proximity to each other ; setae Mr are strongly plumose. The extra-marginal setae are plumose and increase in length towards the posterior end of the idiosoma. The lacinae of the tritosternum are long and pilose. The sternal shield is ornamented with strong ridges and tubercles ; the three pairs of sternal setae are simple. The elongate metasternals each carry a simple seta. The genital shield is ornamented with punctate lines; the genital setae are simple. The ventri-anal shield is evenly reticulated and tuberculated, and is provided with five pairs of pre-anal setae of which the external posteriors are pilose. The shape of the shield and the length of the setae show considerable variation in the specimens examined (cf. Text-figs. 45 BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH ‘ NK N y \, A ae ee” re 0 9 SRE weve eevee UG - Che oe = ce . wwe” ~eMee ove weve Variation in the chaetotaxy of the dorsal shield of Geholaspis (Longicheles) Fics. 66-67. mandibularis (Berl.), female. 46 BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH 68-70). The metapodals are elongate and weakly sclerotized. The posterior half of the peritrematal shield is free. The external posterior rostrals lie well in advance of the internals and the ventral groove is provided with five transverse rows of denticles. The corniculi are long and slender and extend beyond the middle of the palp femur. The form of the tectum is shown in Text-fig. 71. The chelicerae are very long and strongly toothed (Text-fig. 72). Leg I (approx. 600 in length) with the tibia (approx. rooy.) shorter than the tarsus (approx. 130/1). Dimensions. Female: length, 750-S0on; breadth, 480—500y. HABITAT AND LocALiTy. In litter, moss and compost. Widely distributed in Europe (Franz, 1954). Genus MACRHOLASPIS Oudemans. Macrholaspis Oudemans, A. C. (1931). Ent. Ber. 8, No. 180: 272. This genus was proposed by Oudemans (1931) for Gamasus opacus C. L. Koch and was characterized by the female having only two pairs of setae on the ventri-anal shield (‘‘ Ventrianaalschild mit 2 paar borstels’’). According to Oudemans one of these two pairs of setae would be the para-anals and the other the pre-anals. Recently, the writers have examined a preparation and an unpublished drawing of opacus from the Oudemans Collection at Leiden, and found that the ventri-anal shield of the female, the only sex known, has two pairs of pre-anal setae and not one pair as Oudemans has stated. Further, Nothrholaspis aciculatus Berl., a common European species, was found to be synonymous with Macrholaspis opacus (Koch) Oudemans, 1931. The following is an emended definition of the genus Macrholaspis : Dorsal shield strongly attenuated posteriorly and bearing twenty-nine pairs! of plumose setae. Lateral margins of the shield markedly serrated. Sternal shield with three pairs of setae ; metasternals free. Genital shield with accessory sclerites and a pair of genital setae. Ventri-anal shield with fwo pairs of pre-anal setae. Other characters as in Macvocheles. Type. Gamasus opacus C. L. Koch, 1839. Two species are represented in the British fauna. these may be separated as follows : 1. Dorsal shield with 58 setae; Mr short and straight (Text-fig. 73). Serrations of the lateral margin of the shield small and rounded (Text-fig. 74). Sternal shield with numerous punctures. Posterior margin of trochanter IV smooth Macrholaspis opacus (C. L. Koch) —. Dorsal shield with 59 setae; Mr long and curved distally (Text-fig. 78). Serrations of the lateral margin of the shield considerably longer and tapering (Text-fig. 80). Sternal shield with a network of ridges and punctures. Trochanter IV with a strong tubercle on its posterior margin 3 “ F ; . Macrholaspis dentatus sp. n. 1 An additional seta may be present in some species (cf. Macrholaspis dentatus sp. N.). = ame are BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH 7\ Bics. 68-72. Geholaspis (Longicheles) mandibularis (Berl.), female. in the form of the ventri-anal shield. Fig. 71, tectum. Fig. Figs. 68-70, variation 72, chelicera. 47 48 BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH Macrholaspis opacus (C. L. Koch) Gamasus opacus Koch, C. L. (1839). Deutsch. Crust. Myr. Arach. fasc. 25, t. 24. Macrocheles (Nothrholaspis) aciculatus Berlese, A. (1918). Redia 13 : 169 syn. nov. Holostaspis terreus, Halbert, J. N. (1915). Proc. Roy. Ivish Acad. 31, ii: 66. Macrholaspis opacus, Oudemans, A. C. (1931). Ent. Ber. 8, No. 180: 272. Nothrholaspis aciculatus, Willmann, C. (1939). Ark. Zool. 31 A, 10: 6, figs. Macrocheles (Nothrholaspis) terreus, Cooreman, J. Bull. Mus. voy. Hist. nat. Belg. 19, 63 : 21. FEMALE. Dorsal shield strongly attenuated posteriorly (Text-fig. 13) and charac- teristically ornamented in its anterior third with a polygonal network of minute spines (aciculatus !). The lateral margins of the shield are distinctly serrated (Text-fig. 74). The vertical setae are strongly developed ; setae M1 are short and plumose (Text-fig. 75). The remaining twenty-seven pairs of plumose setae are distributed as in Text- fig. 73. The extra-marginal setae are of the same form as the marginal series. The tritosternum is normal with the lacinae pilose. The sternal shield is densely covered with punctures and bears three pairs of simple setae (Pl. 4, fig. 18). The metasternal setae are also simple. The genital shield is covered with punctures which tend to form a network in its anterior two-thirds. Accessory sclerites are well developed and the genital setae are plumose. The ventri-anal shield (approximately 220 long and 198m wide) is oval in outline and, like the dorsal shield, ornamented with spinules. The two pairs of pre-anal setae are plumose, the para-anals long and simple, and the post-anal seta strongly plumose. The interscutal membrane between the genital and ventri-anal is provided with three pairs of platelets, one pair of pore- bearing platelets and a pair of plumose setae. The metapodal shields are conspicuous. The peritreme and peritrematal shield are normal. The gnathosoma bears four pairs of setae ventrally. The ventral groove has five transverse rows of denticles. The form of the tectum is shown in Text-fig. 76. Both digits of the chelicera are bi-dentate ; the distal tooth on the fixed digit is small and difficult to see if the chelicera is not orientated correctly (Text-fig. 77). The pilus dentilis is long and stout. The dorsal seta is comb-like. Leg I (530) has plumose setae on the femur, genu and tibia. Tibia I (85yz) is con- siderably shorter than the tarsus (121j). Legs II to IV measure 550, 495 and 800, respectively. Mare. Unknown. Dimensions. Female; length 725-740” ; breadth 445-470. HABITAT AND LocALITy. Widely distributed in Europe. Common is decaying wood, moss and litter. Macrholaspis dentatus sp. n. FEMALE. Dorsal shield attenuated posteriorly as in the preceeding species but without spinules. The ornamentation consists of a network of lines and punctures (Text-fig. 78). The chaetotaxy of the shield comprises fifty-nine setae arranged as in the figure. The verticals are stout and setae Mr long, plumose and curved (Text-fig. 79). An interesting feature of the chaetotactic pattern is the presence of three : BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH 49 7, 76 Fics. 73-77. Macrholaspis opacus (C. L. Koch), female. Fig. 73, dorsal shield. F. ig. 74, lateral margin of dorsal shield. Fig. 75, anterior region of dorsal shield. Fig. 76, tectum. Fig. 77, chelicera. ZOOL, 4, I. 50 BRITISH MITES-SUBFAMILY MACROCHELINAE TRAGARDH unpaired setae in the dorsal series. The lateral margins of the shield are strongly serrated (Text-fig. 80). The extra-marginal setae are similar to the marginals. The sternal shield is provided with a network of ridges and punctures and bears three pairs of simple setae (PI. 4, fig. 19). The metasternal shields are small and free, and the setae simple. The genital shield is strongly ornamented with a network of ridges and punctures ; the genital setae are plumose. The ventri-anal shield, similarly 80 Fics. 78-80. Macrholaspis dentatus sp. nov., female. Fig. 78, dorsal shield. Fig. 79, anterior region of dorsal shield. Fig. 80, lateral margin of dorsal shield. ornamented, measures 187y in length and 145 in breadth, and bears two pairs of plumose pre-anal setae. The para-anals are long and simple, and the post-anal seta short and plumose. The interscutal membrane between the genital and ventri-anal shields is provided with platelets and plumose setae as in the preceding species. The peritreme and peritrematal shield are normal. The chaetotaxy and structure of the gnathosoma are essentially the same as in opacus. The form of the tectum and the dentition of the chelicerae are also similar. Leg I, approximately 525y in length, has the tibia considerably shorter than the BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH 51 tarsus (tibia, 88; tarsus, 1104). Legs II to IV measure 575, 560” and 88oy in length, respectively. Dimensions. Length 7654; breadth 430. HABITAT AND Locality. A single female from humus under bracken in the Leri Valley, North Cardiganshire, Wales. Holotype female, 1955.10.22.95. This species is characterized by the form of setae M1, the ornamentation of the sternal shield and the shape of the ventri-anal shield. Genus HOLOSTASPELLA Berl. Holostaspella Berlese A. (1904). Redia 1: 241 Dorsal shield with twenty-eight pairs of setae. Vertical setae situated on an outgrowth of the dorsal shield. Sternal shield with three pairs of setae ; metasternal shields free. Ventri-anal shield with four pairs of pre-anal setae. Gnathosoma normal. Leg I in the female armed with stout spurs. Other characters as in Macrocheles Latr. Type. AHolostaspis (Holostaspella) sculpta Berl., 1904. This genus is represented in Britain by one species only. Holostaspella ornata (Berl.) Macrocheles vagabundus, Oudemans, A. C. (1902). Tijdschr. Ent. 45: 43, figs. Holostaspis ornatus Berlese, A. (1904). Redia 1: 277. Holostaspella ornata Oudemans, A. C. (1931). Ent. Ber. 8 : 273, syn. nov. Holostaspella ovnata, Evans, G. O. (1956). Proc. Linn. Soc. Lond. FEMALE. Dorsal shield, finely reticulated, bears twenty-eight pairs of setae (Text-fig. 81). The verticals are stout and pilose, and are situated on a prominent projection (Text-fig. 82). The setae on the shield are minutely pilose as shown in the figure. The sternal shield is massive and characteristically ornamented with ridges and depressions (PI. 4, fig. 20). The first pair of sternal setae are strongly plumose ; the second and third pair smooth. The metasternals, triangular in outline, each bear a simple seta. The large genital shield is richly ornamented ; the genital setae simple. Most of the region posterior to coxae IV is occupied by the heavily ornamented ventri-anal shield (335 x 315/), which bears four pairs of pre-anal setae in addition to the para-anals and the post-anal seta. The anterior pre-anal is weakly pilose and the post-anal plumose. The interscutal membrane is coarsely striated. The gnathosoma is normal for the Macrochelinae. The lateral processes of the tectum are free (Text-fig. 83). The dentition of the chelicera is shown in Text-fig. 84. Leg I with the tibia (121) shorter than the tarsus (132). The femur and trochanter of leg II are spurred ; the tarsus has a large stout spine ventrally (Text-fig. 85). Mare. Unknown. 52 BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH 82 Fics. 81-85. Holostaspella ornata (Berl.), female. Fig. 81, dorsal shield. Fig. 82, anterior region of dorsal shield. Fig. 83, tectum, Fig. 84, chelicera. Fig. 85, leg II (ambulacrum omitted). oa eet BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH 53 Dimensions. Length 924-950; breadth 550-560y. HABITAT AND LOCALITY. This species has previously been recorded from the Netherlands (Oudemans, 1902) and Austria (Franz, 1954). The writers have examined a single specimen taken from Sphaerocerus sp. captured at Bagley Wood, Berks (Coll. O. W. Richards, det. Vitzthum). The above description and figures are based on the type from the Oudemans Collection, Leiden. SUMMARY 1. The classification of the Macrochelidae is discussed and keys are given for the identification of British species of the Macrochelinae. 2. The following three new species are described and figured : Macrocheles rothamstedensis sp. nov. Macrocheles punctoscutatus sp. nov. Macrholaspis dentatus sp. nov. 3. The following species are recorded for the first time from Britain : Macrocheles carinatus (C. L. Koch) Macrocheles decoloratus (C. L. Koch) Macrocheles insignitus (Berl.) Macrocheles montanus (Willmann) Macrocheles penicilliger (Berl.) Macrocheles subbadius (Berl.) Macrocheles tardus (C. L. Koch). Geholaspis (Longicheles) mandibularis (Berl.) Holostaspella ornata (Berl.) 4. The following species have been relegated to the synonymy : Nothrholaspis aciculatus Berl., 1918 syn. of Macrholaspis opacus (C. L. Koch) 1839. Holostaspella ornata Oudemans, 1931, syn. of Holostaspella ornata (Berl.), 1904. Macrocheles gladiator Hull, 1918, syn.(?) of Macrocheles plumiventris Hull, 1918. Macrocheles plumipes Hull, 1918, syn.(?) of Macrocheles plumiventris Hull, 1918. Macrocheles minimus Hull, 1918, syn. of G. (Longicheles) mandibularis (Berl.), 1904. Macrocheles hulli Falconer, 1923, syn. of Macrocheles carinatus (C. L. Koch), 1839. Macrocheles occidentalis Hull, 1925, syn. of Macrocheles submotus Falconer, 1924. Macrocheles gloriosus Hull, 1925, syn. of Macrocheles submotus Falconer, 1924. Nothrholaspis fimicola Sellnick, 1932, syn. of Macrocheles plumiventris Hull, 1925. Coprholaspis anglicus Turk, 1946, syn. of Macrocheles glaber (Miiller), 1860. 54 BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH ACKNOWLEDGMENTS We are extremely grateful to Dr. L. van der Hammen for the loan of specimens from the Oudemans Collection at Leiden ; to Dr. F. A. Turk for the loan of the type of Coprholaspis anglicus Turk; to Dr. G. Lombardini for comparing some of our specimens with the types in the Berlese Collection, Florence and to Mr. M. G. Sawyers, British Museum (Nat. Hist.) for the photomicrographs. REFERENCES BERLESE, A. 1882. Gamasidi nuovie poco noti. Boll. Soc. ent. ital. 14 : 338-352. 1882-1903. Acari, Myrviopoda et Scorpiones hucusque in Italia veperta, Portici et Padua. Fasc. I—101. 1904. Acarinuoviland II. Redia, 1 : 235-280. 1918. Centuria quatria di Acari nuovi. fFedia, 13: 115-102. 1921. Indice sinonimico dei generi e delle specie illustrate nei fascicolo 1 a ror. Redia, 14 : 77-105. CoorEMAN, J. 1943. Note sur la faune des Hautes-Fagnes en Belgique (1) XI. Acariens (Parasitiformes) (2). Bull. Mus. voy. Hist. nat. Belg. 19, No. 63 : 1-28. Evans, G. O. 1956. An introduction to the British Mesostigmata with keys to familes and genera, J. Linn. Soc. Lond. Zool. (in press) Fatconer, W. 1923. Two British mites new to science and a new subgenus of Macrocheles Latr. Naturalist, Lond. : 151-153. 1924. Macrocheles submotus—a new name for M. cognatus Falcr. (nom. praeocc.). Naturalist, Lond. : 363. Franz, H. 1954. Die Nordost-Alpen. 15. Ordnung Acarina in Spiegel Land-Tvierwelt Inns- bruck, 1 : 329-452. HERMANN, J. F. 1804. Mémoire aptévologique. Strasbourg, pp. vi + 144, 9 pls. Havsert, J. N. 1915. Clare Island Survey. Acarinida: ii. Terrestrial and marine Acarina. Proc. voy. Ivish Acad. 3 39 i1: 45-136. Hucues, A.M. 10948 . Mites associated with stored food products. London: H.M.S.O., pp. 168. Hutt, J. E. 1918. Terrestrial Acari of the Tyne Province. Tvans. nat. Hist. Soc. Northumb. N.S., 5: 13-88. — 1925. Acari of the family Gamasidae: new and rare British species. Ann. Mag. nat. Hist. (9) 15 : 201-219. Hyatt, K.H. 1956. A collection of mites from stable manure. Ext. mon. Mag. (in press). Kocu, C. L. 1836-1841. Deutschlands Crustaceen, Myviopoden und Avachniden, Regensburg. Kramer, P. 1876. Zur Naturgeschichte einiger Gattungen aus den Familie der Gamasiden. Arch. Naturgesch. 42 : 46-105. MULLER, J. 1860. Insectenepizoen der mahrischen Fauna (Jahr. naturw. sect.) K.K. mahr. schles. Gesell. Ackerbau, Natur.-u. Landes : 157-184. Oupemans, A. C. 1901. Bemerkungen tiber Sanremeser Acari. Tijdschr. Ent. 43 : 129-139. 1913. Acarologische Aanteekeningen. Ent. Ber. 4, 73 : 2-18. 1914. Acarologisches aus Maulwurfsnestern. Aych. Naturgesch. 79a, 8 : 108-200. 1931. Acarologische Aanteekeningen CIX. Ent. Ber. 8, 180 : 272-280, PeREIRA, C. & Castro, M. P. DE. 1945. Contribuicao para o conhecimento da especie tipo de Macrocheles Latr. (Acarina): M. muscaedomesticae (Scopoli, 1772) emend. Arq. Inst. Biol. S. Paulo, 16 : 153-186. Scopoul, J. A. 1772. Annus V. Historico naturalis. Lipsiae. BRITISH MITES—SUBFAMILY MACROCHELINAE TRAGARDH 55 SELLNICK, M. 10931. Zoologische Forschungreise nach den Jonischen Inseln und dem Peloponnes von Max Beier, Wien. Acari. S.B. A kad. Wiss. Wien, 140: 693-776. 1940. Die Milbenfauna Islands. Goteborg. Vetensk. Samh. Handl. (5) 68, 14: I-129. TrAGARDH,I. 1949. Description of two new genera of Mesostigmata (Acarina). A spidilaelaps from Samoa and Protoholaspis from Peru. Ent. Medd. 25 : 311-325. 1952. carina collected by the Mangarevan expedition to South Eastern Polynesia in 1934 by the Bernice P. Bishop Museum, Honolulu, Hawaii. Mesostigmata. Ark. Zool. (2) 4: 45-90. Turk, F. A. 1946. Studies on Acari. V: Notes on and descriptions of new and little- known British Acari. Ann. Mag. nat. Hist. (t1) 12 : 785-820. 1948. Insecticolous Acari from Trinidad, B.W.I. Proc. Zool. Soc. Lond. 118 : 82-125. VaLLE, A. 1953. Revisione di generi e sottogeneri berlesiana di Acari. Redia, 38: 316-360. WILIMANN, C. 1939. Die Arthropodenfauna von Madeira... etc. XIV. Terrestrische Acari (exkl. Ixodidae). Avk . Zool. 31a 10: I-42. —— 1951. Die hochalpine Milbenfauna der mittleren Hohen Tauern insbesondere des Grossglockner Gebietes. (Acari). Bonney Zool. Beity.2: 141-176. 28 APR 3b PRESEN EXPLANATION OF PLATES PLATE 1 Sternal, genital and ventri-anal shields of the females of : Fic. Fic. Fic. Fic. Fic. Fic. Ie CE aS Macrocheles muscaedomesticae (Scopoli). 95. Macrocheles vothamstedensis sp. nov. X 153- Macrocheles glabey (Miiller). 115. Macrocheles punctoscutatus sp. nov. X 110. Macrocheles subbadius (Berl.). x 160. Macrocheles insignitus Berl. X 218. Bull. B.M. (N.H.) Zool Anes LUN ANID, Sternal, genital and ventri-anal shields of the females of : Fic. 7. Macrocheles mervdarvius (Berl.), 199. Fic. 8. Macrocheles montanus (Willmann). » 100. Fie. 9. Macrocheles cavinatus (C. L. Koch). roo. Fic. 10. Macrocheles penicilliger (Berl.). 110. Fic. 11. Macrocheles submotus Falconer. ™ 90. Fic. 12. Macrocheles tardus (C. L. Koch). Ito. Bull. B.M. (N.H.) Zool. 4,1 PLATE 3 Sternal, genital and ventri-anal shields of the females of : Fic. 13. Macrocheles decoloratus (C. L. Koch). x 120. Fic. 14. Macrocheles matrius Hull. 112. Fic. 15. Macrocheles plumiventris Hull. x 74. Fic. 16. Macrocheles superbus Hull. 70. 11 16; on 4 IPT JAN Bull. B.M. (N.H.) Zool. 4, 1. PLATE 4 Sternal, genital and ventri-anal shields of the females of : Fie. Fic. Fic. Fic. 1s 18. 19. 20. Geholaspis (Geholaspis) longispinosus (IXramer). Macrholaspis ofacus (C. L. Koch). 153. Macrholaspis dentaius sp.nov. X 131. Holostasfella ovnata (Berl.). * 123. x 100. Bull. B.M. (N.H.) Zool. 4, 1. :VOLUTION OF _ RATITES + - : erat SIR GAVIN de BEER BULLETIN ‘OF THE EVOLUTION OF RATITES BY SIR _GAVIN de BEER, F.R.S. Pp. 57-70, Plates 5~9 Gh 7 s& owe » ae "e% , BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 4 No. 2 LONDON: 1956 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, 1s issued in five series corresponding to the Departments of the Museum, and an Historical Series. Parts appear at irregular intervals as they become veady. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. This paper is Vol. 4, No. 2 of the Zoological series. PRINTED BY ORDER OF THE TRUSTEES OF THE BRITISH MUSEUM Issued August, 1950 Price Ten Shillings THE EVOLUMON, OF RATITES By Sir GAVIN bE BEER, F.R.S. Director, British Museum (Natural History). SYNOPSIS Adequate knowledge of the structure of Archaeopteryx now enables a comparison to be made between it and the Carinates. In the latter the structure of the wing, the tail, and the cerebellum can be shown to be adaptations to flight. Since these same adaptations are found in Ratites, they would be inexplicable unless the Ratites were descended from flying birds. The palate of the Ratites is not primitive but neotenous, and represents an early stage through which the palate of many Carinates passes during development. Other neotenous features of the Ratites are the plumage and the persistence of sutures between the bones of the skull. INTRODUCTION In the first edition of the Origin of Species (1859, p. 134) Darwin wrote: “‘ As Professor Owen has remarked, there is no greater anomaly of nature than a bird that cannot fly; yet there are several in this state.’’ There was a touch of irony in making this quotation, for as is well known, Owen’s views on evolution were uncertain and equivocal, and the very existence of flightless birds was inescapable evidence of descent with modification from the “archetype” of birds. Darwin himself continued: “‘ We may believe that the progenitor of the ostrich genus had habits like those of the bustard, and that, as the size and weight of its body were increased during successive generations, its legs were used more, and its wings less, until they became incapable of flight.” A few years later, in his treatise on The Anatomy of Vertebrates (1866, 2, 12.) Owen put forward the view that the Cursores or Ratites were not “‘ a natural order ; some of its exponents have demonstrably closer affinities to other groups of which they are wingless members.’’ Further on (Joc. cit. : 43), Owen referred to the Ratites as “those birds in which the power of flight is abrogated’. For a man who did not believe unreservedly in evolution, this was about as near as he could get to the view that the Ratites are descended from flying birds, and he even supplied the explanation of such a descent (loc. cit.: 12): ‘‘ by the arrested development of the wings unfitting them for flight ’’. The view of the degenerate nature of the Ratites has been supported by M. Fiirbringer (1888), T. J. Parker (1892), W. P. Pycraft (1901), R. Broom (1906), J. E. Duerden (1920), E. Stresemann (1927-34), W. K. Gregory (1935), and many others ; and it might have been thought that the evolution of flightless birds from flying birds was generally accepted. Nevertheless the hypothesis has been put forward that the structure of the Ratites is in many respects so primitive that they must have ZOOL, 4, NO. 2. 5 60 THE EVOLUTION OF RATITES branched off from the main stem of bird evolution before the power of flight was acquired. B. Lindsay (1885), R. S. Wray (1887), A. C. Chandler (1916) and J. C. Ewart (1921) are of this opinion, but the foremost exponent of this view has been P. R. Lowe (1928, 1935), with whom M. Friant (1945, 1946) has expressed agreement. It has been known for a long time that the Ratites show a number of characters which have been considered as primitive. Among these is the palate, on which T. H. Huxley (1867) based a system of classification of the birds in which the dromaeogna- thous type, characteristic of the Ratites, was regarded as the most primitive. Pycraft (1900, Igor) who extended Huxley’s observations, summed up (1901 : 343) the situation as he saw it in the words: “ The contention that the Struthious (Palaeognathine) palate is of a more ancient type than the Neognathine is admitted by all.’ The term palaeognathine or palaeognathous is equivalent to dromaeo- gnathous, while neognathine or neognathous includes all the other categories of Huxley’s classification. This argument has been adopted by Lowe, in whose view the Ratites show “ the primitive palate from which the neognathous palate characteristic of the modern or flying birds was obviously derived’”’. To this Lowe has added six further argu- ments, all in support of his view that the ancestors of Ratites never flew. They are : 1. The “ primitive ’’ disposition of the muscles. 2. The fact that “‘ all the feathers borne by the adult ostrich or by any other struthious form whether they are situated on the wing, or body generally, are nothing more than down, or modified down,” and being juvenile structures are he thinks therefore ancestral and primitive. 3. The absence of the rudiment of the clavicle in the embryo of the ostrich, which, in his view, is proof that the ostrich decended from ancestors which had lost the clavicle, and therefore not from flying birds in which the clavicle is preserved. 4. The persistence of the sutures in the skull of the ostrich, in which it resembles the condition of the ancestral reptiles and differs from that of flying birds in which the bones of the skull are firmly fused in the adult. 5. The obtuse angle subtended between the coracoid and the scapula, which resembles the condition of the ancestral reptiles and differs from that of flying birds where this angle is more or less acute. 6. The similarity between the bones of the hand of the ostrich and those of a dinosaur such as Oynitholestes. With perfect logic, Lowe contended that if, as he believed, the Ratites were descended from birds which had never acquired the power of flight, then it must follow that Avchaeopteryx could not have been in the line of ancestry of birds, but must have been an independent offshoot from the reptiles. The detailed knowledge now available of the structure of Avchaeopteryx (de Beer, 1954) can be used to test this hypothesis. Two related problems are involved : the evolution from reptiles of birds in general, and the evolution of Ratites in particular. These problems can be solved by finding the answers to the following three questions : THE EVOLUTION OF RATITES 61 1. Is Archaeopteryx on the line of evolution from reptiles to birds? 2. Arethere any characters by which modern flying birds differ from Archaeopteryx which can be attributed with certainty to adaption to active flight? 3. Are these characters also shown by the Ratites? ARCHAEOPTERYX AND THE ANCESTRY OF BIRDS If Archaeopteryx was the product of an independent line of evolution from the reptiles, unrelated to the stock which gave rise to birds, it then becomes necessary to believe that the feathers of Archaeopteryx and the feathers of all other birds were independently evolved. The identical details of structure which the feathers show involve the quill, the vane formed of barbs, held together and parallel with one another and yet capable of being torn apart, the proportions between the vane and the quill and between the proximal and distal portions of the vane. In all these respects, the structure of the feather in Archaeopteryx and in modern flying birds is so exactly identical that it is impossible to believe that they were independently evolved. But this is not all. In addition to the feathers themselves, there is the manner in which they are arranged on the wing, the differentiation between larger feathers or Temiges and smaller feathers or coverts, and the further differentiation of the remiges into primaries, borne on the wrist-joint and hand, and secondaries borne on the forearm. Here again the conditions are identical in Archaeopteryx and in modern flying birds. It follows that the view that Archaeopteryx is not related to the modern birds is completely untenable. Granting that Archaeopteryx represents an example of an early stage in the evolu- tion of feathered organisms away from the reptiles, it may still be asked whether Archaeopteryx is ancestral to modern birds. The remarkable mosaic of reptilian and avian characters that Archaeopteryx shows has been discussed elsewhere (de Beer, 19542). The conclusion to be drawn is that Archaeopteryx is a vara avis among fossils in that it is possible to say that nothing is known, either by way of structures which it possesses or does not possess, or of the time-relations of its occurrence, which might disqualify it from being regarded as a true ancestor of modern birds. AsG.G, Simpson (1936: 92) has said, “ every difference between Archaeopteryx ... on one side and true reptiles of possible ancestral type, especially the Pseudosuchia, on the other, is definitely in the direction of true birds”. If, as H. Steiner (1918, 1956) believes, Archaeopteryx was aquintocubital, it would provide yet another proof that it was ancestral to modern birds. CARINATES AND ADAPTATIONS TO FLIGHT Accepting the fact that Archaeopteryx is a mile-stone on the road from reptiles to modern birds and represents the type of structure from which Carinates evolved, attention may be turned to the question whether any of the differences observable between Archaeopteryx and the modern flying birds or Carinates can with certainty be ascribed to adaptation to flight. That the flying bird is highly adapted to its 62 THE EVOLUTION OF RATITES mode of life is a commonplace of biological expression, and in the case of some structures it is easy to prove it. Attention will here be confined to the carpo- metacarpus, the pygostyle, and the cerebellum. The keel on the sternum is deliberately omitted from the discussion since its absence in the Ratites is the basis of their diagnosis, and the question at issue is whether this absence is primitive or specialized. The carpometacarpus is the product of fusion between the distal carpals and the three metacarpals, the 2nd and 3rd of which are fused again at their distal extremities. The result is a structure providing a light and resilient yet firm basis for the attach- ment of the primary remiges. It is absolutely characteristic of modern birds and found nowhere else. In Archaeopteryx the forelimb skeleton consists of proximal carpals, the radiale and ulnare which remain more or less separate, distal carpals fused together and to the base of the third metacarpal, and separate and independent 1st and 2nd metacarpals. Archaeopteryx was unable to do much more than glide, and as Dr. H. W. Parker has remarked to me, the air-pressure on the feathers of its wings must have been lower than in an actively flying bird with the same ratio of wing-area to mass, because Archaeopteryx was unable to maintain itself in the air continuously against the pull of gravity. The carpometacarpus of the Carinate is without doubt an adaptation to flight by enabling the wing to exert and withstand greater pressure. There is one further feature of the wing of Carinates that calls for notice, and that is the presence of a small number of feathers attached to the first digit of the hand, forming a “‘ bastard wing’’. These few feathers add nothing to the weight-bearing power of the wing, yet they perform a function of capital importance in flight for they serve like the slotted wing of a modern aircraft to maintain a slip-stream of air and prevent stalling. The “ bastard wing ’”’ is a beautiful adaptation to flight. In Archaeopteryx the tail is very long, as long as the rest of the body, and its skeleton consists of 20 elongated separate vertebrae, to each of the hinder 15 of which correspond a pair of rectrices, quill-feathers, which form an oblong and elongated air-resisting surface. In Carinates the tail is very short, consisting of about a dozen flattened vertebrae, the hindmost half-dozen or so of which are fused together, giving rise toa pygostyle. The rectrices, to the number of a dozen pairs, are disposed transversely. A masterly analysis and comparison of the conditions in the tail of Archaeopteryx and Carinates has been given by H. Steiner (1938). It has been pointed out by J. Maynard Smith (1952) that the structure of the most primitive flying animals is one that imparts aerodynamic stability. That is to say that they are of a shape such that when in “ flight ’’ through the air, they are able to maintain the direction of their progress without muscular intervention and compensatory movements. In other words, such animals are gliders, and the structure of the skeleton, wings, and tail of Avchaeopteryx is just such as would have enabled it to glide with stability, but not fly actively. The perfection of the power to fly has involved the development of the ability to perform mechanically unstable flight-movements, such as rapid pitching, yawing, and banking, for which a reduction of the long axis of the animal is essential. The pygostyle of the Carinates is without doubt an adaptation to flight, THE EVOLUTION OF RATITES 63 The cerebellum of Archaeopteryx is best characterized negatively by saying that it is small and does not overlap forwards over the midbrain. In other words, the brain of Archaeopteryx is similar to that of reptiles. In Carinates, on the other hand, the cerebellum is so large that it expands forwards as a median and unpaired structure over the dorsal surface of the midbrain, which it presses downwards, and so the cerebellum comes into contact with the hinder part of the cerebral hemisphere. The result is that the cerebellum of Carinates hides the optic lobes, whereas the latter structures are plainly visible in Archaeopteryx. The cerebellum has been defined by Sherrington as the head-ganglion of the proprioceptive system. As L. Edinger (1912: 300) has shown in Columba, among the most important sources of impulses conducted to the cerebellum are the organs of balance in the semi-circular canals of the ear, which respond not only to changes in static conditions, but also to changes in the dynamic conditions of the organism caused by alterations in speed and direction of motion. In the case of Carinates, as flying birds the possibilities of direction of motion are greatly increased by the introduction of the vertical dimension. At the same time, the performance of flight requires high speed of adjustment and compensatory movements, not only in actual flight but in landing on small objects. As J. Z. Young (1950 : 455) has said, there is reason to think that the large size of the cerebellum in flying birds is connected with the precise control of movement in all planes of space during flight. The view that the cerebellum of Carinates is an adaptation to flight is confirmed by the conditions in the pterosaurs. There, as T. Edinger (1941 : 678) has shown, there is ‘“‘ a cerebellum thrust forward above the midbrain to adjoin the forebrain as in birds ; obviously this is one of the characters distinguishing all pterosauria from the other reptiles”’. There can be no doubt that the parallel development and large size of the cerebellum in pterosaurs and in birds, by which they both differ from all non-pterosaurian reptiles, are due to the same cause: adaptation to flight. Having now established that the carpometacarpus, the “‘ bastard wing’’, the pygostyle, and the large size of the cerebellum of Carinates are adaptations to flight, attention may be turned to the conditions in Ratites in respect of these structures. ADAPTATIONS TO FLIGHT IN RATITES The skeleton of the wing of Ratites is built on identically the same plan as that of the Carinates. B. W. Tucker (1938a: 224) has stressed the similarity not only in the points of fusion between the various elements which go to make up the carpometa- carpus in both Carinates and Ratites, but also subtle points, such as the curvatures of the 2nd and 3rd metacarpals and the expansion of the basal phalanx of the 3rd digit. H. Steiner (1949 : 367) has studied the wings of Carinates and Ratites by means of X-rays and concludes that it “‘lasst sich ohne jeden Zweifel feststellen, dass genau die gleichen Eigentiimlichkeiten zu beachten sind. Ausgehend von irgend einem Carinatenfliigel kann ausserdem tiber den Fliigel von Rhea und Struthio bis zu jenem von Casuarius eine zunehmende Verkiimmerung verfolgt werden, welche iiber die Zustande, wie sie bei Dromaeus und Apteryx angetroffen werden, bis zur vollstandigen 64 THE EVOLUTION OF RATITES Reduktion des Fliigels bei den ausgestorbenen Riesenstraussen Aepyornis und Dinornis gefiihrt hat ”’. There can be no doubt that the skeleton of the wing in the Ratites shows features associated with adaptation to flight which are explicable only on the view that they were inherited from ancestors which flew. The skeleton of the tail in Ratites has been studied by W. Marshall (1873) and referred to by W. K. Gregory (1935), but otherwise has not attracted much attention. It is composed of a varying small number of vertebrae which in some forms decrease in size caudally and taper out. But in the ostrich there is a structure composed of the fusion of the terminal vertebrae which undoubtedly constitutes a pygostyle. It is Shown in PI. 5 in comparison with a Carinate pygostyle. Since this structure in Carinates is certainly associated with the power of flight, its presence in a Ratite is inexplicable unless the ancestors of Ratites also flew. This has also been pointed out by Gregory. The cerebellum of birds has been subject to an exhaustive study by S. Ingvar (1918), the results of which show that the large size of the cerebellum in Carinates is matched by a similar large size in Ratites. Not only does the cerebellum of the ostrich, for instance, project forwards over the dorsal surface of the mid-brain towards the cerebral hemisphere, but it shows the same arbor vitae structure as the cerebellum of a Carinate when seen in sagittal section. In Plate 6 are shown the brain of Archaeopteryx in side view, and sagittal sections through the brains of Rhea and Tringa. It is clear that the structure of the cerebellum is the same in the Ratites as in the Carinates ; and if its structure in the latter is an adaptation to flight, its structure in the former is inexplicable unless the Ratites were descended from flying birds. On all three counts, the evidence is conclusive that the Ratites must have evolved from flying birds. It remains now to consider a few further points which receive ready explanation on this view, and to refute the grounds on which Lowe thought that the Ratites were primitively flightless. It has been shown above that the skeleton of the wing of the Ratites bears evidence of adaptation to flight. It may be added that in one form, the Rhea, there is still to be seen a trace of the differentiation between primary and secondary remiges, as shown in Plate 7. This distinction, which already exists in Archaeopteryx would be meaningless unless the Rhea’s ancestors had been capable of flight. Even more remarkable is the presence in the Rhea of feathers on the 1st digit forming a ‘“‘ bastard wing ”’, an adaptation evolved in Carinates which results in the maintenance of the slip-stream in flight. The curious phenomenon of diastataxy or aquintocubitalism, the absence of the 5th secondary remex from the row of flight feathers in the wing, has long been a puzzle. Its most probable explanation has been provided by H. Steiner (1918) who has shown in a brilliant and exhaustive series of studies that it is associated with the peculiar method of folding the wing, the ulnar flexure, adopted by birds. When a bird folds its wings, the hand is moved sideways relatively to the forearm through an angle of almost 180°. The development of this new type of movement affected the feather-rudiments in the skin at the point of flexure and dislocated them in such a way that the rudiment which would have given rise to the 5th secondary remex is THE EVOLUTION OF RATITES 65 displaced, and, instead, develops into the 5th major covert, leaving a gap in the series of secondaries. Steiner has shown conclusively that in the Carinates the aquintocubital condition is primitive, and that the presence of the 5th secondary remex, which is found sporadically in some members of nearly all groups of birds, is due toasecondary readjustment. Be that asit may, it is clear that the phenomenon of aquintocubitalism is intimately associated with the structure and arrangement of the remiges in a flying wing. It is therefore remarkable that a vestige of the aquinto- cubital condition is found in the wing of the young Apteryx (Steiner 1918 : 434) which thereby is shown to possess a structure characteristic of primitive Carinates and which could not have been independently evolved. Professor Steiner has kindly informed me that he has evidence that other Ratites also are aquintocubital. Further, there is another line of evidence relating to the loss of the power of flight of Apteryx. R. Broom (1947 : 49) has ingeniously shown that as New Zealand has had no land connexions with any other continent since early Jurassic times, and as the centre of evolution of birds exemplified by Archaeopteryx was situated in the Palaearctic continent in middle Jurassic times, the ancestors of Apteryx could not have reached New Zealand unless they flew thither. THE NEOTENY OF RATITES Reverting now to the reasons on which Lowe sought to base the view that the Ratites were primitive birds whose ancestors had never flown, one: the similarity between the hand of the ostrich and that of the dinosaur, has been dismissed as invalid. Tucker (1938) has shown that such resemblances as there are between them are only superficial and without significance. Another: the angle between the coracoid and the scapula, can be shown to be due to the reduction of the length of the pectoral muscles in the Ratites ; for it is the lengthening of the coracoid in the Carinates which is responsible for the acuteness of the angle between the coracoid and the scapula; and the length of the coracoid may be regarded as an adaptation to flight since it is associated with the lengthening of the pectoral muscles. Whether the disposition of the muscles in the Ratites is “ primitive ’’, as Lowe has contended, is a matter for argument ; but what is no matter for argument is the explanation of the presence in the Ratites of nestling-down, permanent sutures between the bones of the skull, and the dromaeognathous structure of the palate. All these are demonstrably the result of neoteny or the secondary retention of features which were juvenile in the ancestors of the Ratites. To begin with the feathers. It is well known that the down-feathers, nestling- down or neossoptyles, are nothing but the fluffed-out distal ends of the rudiments of the adult feathers or teleoptyles. In Carinates, particularly those in which the young are nidifugous and have a “‘ chick” stage, the nestling-down is well developed, and it owes its fluffiness to the fact that the barbs have no hooks and therefore the feathers form no vanes. This nestling-down is subsequently discarded when the adult feathers or teleoptyles take the place of their former distal extremities the neossoptyles. That the Ratites are neotenous in retaining their ‘‘ ostrich feathers ”’ or nestling-down throughout life is admitted by Lowe himself (1935 : 420) : “‘ So far , 66 THE EVOLUTION OF RATITES as their feather covering is concerned the Struthiones are big, overgrown chicks. They are the “‘ Peter Pans’’ of the avian world. They have never grown up.”’ The same phenomenon of neoteny is responsible for the retention of the sutures between the bones of the skull in the ostrich. In the Carinates, the sutures between the bones are present in the young stages, but they are obliterated in the adult skull, which is a structure of great solidity, in all probability adapted to the necessity for withstanding the mechanical stresses consequent upon active flight. In retaining the sutures between the bones of the skull the ostrich, having lost the power of flight, shows a secondary return to the juvenile condition of the ancestral flying bird, and, of course, the ancestral reptile. The inclusion of the dromaeognathous or palaeognathous palate among the neotenous features of the Ratites, with the implication that it is the result of a secondary retention of an ancestral juvenile condition, may appear surprising in view of the selection of this very structure by T. H. Huxley as the basis for his classification of birds, and his view that the dromaeognathous type was primitive. Nevertheless, the evidence is quite clear, as W. P. Pycraft (1900, rgor) has shown, although he did not realize its significance. The so-called palaeognathous palate is an arrested stage in the development of the neognathous palate. Precisely the same conclusion was reached by S. McDowell (1948) on other grounds, namely the impos- sibility of giving a definition of the palaeognathous palate applicable to all Ratites and tinamus and excluding all Carinates (except tinamus) because of its great variation. The essential feature of Huxley’s dromaeognathous and Pycraft’s paleognathous palate is the fact that the pterygoids extend forwards and make contact with the hinder ends of the prevomers, while the palatines lie further to the side. In Huxley’s schizognathous and aegithognathous types, or Pycraft’s neognathous palates, the usual condition in the adult is that the pterygoids do not make contact with the prevomers, but are separated from them by the palatines with which the pterygoids make a joint. But Pycraft’s remarkable discovery, to which insufficient attention has been paid, was that in the development of many “neognathous”’ birds the palate passes through a “‘ palaeognathous ”’ stage in which the pterygoids actually or nearly come into contact with the prevomers ; but the anterior ends of the pterygoids then become detached from the remainder of these bones, and, instead, become attached to the hinder ends of the palatines, where they give rise to the so-called ‘““mesopterygoid”” elements of W. K. Parker (1875, 1876, 1877, 1879), and the ‘“‘hemipterygoid’’ of Pycraft. Between the detached anterior portion and the remainder of the pterygoid a joint is formed. This is why in the adults of these birds the pterygoid seems not to reach the prevomer, whereas morphologically, in fact, it does or almost does reach it. The hemipterygoid in various Carinates is shown in Plates 8 and g for comparison with the conditions in Ratites. For those, if there be any, who still believe in the theory of recapitulation, it would no doubt be tempting to say that the neognathous palate “‘ recapitulates ” in its development the condition of the palaeognathous palate which would therefore be ancestral. But in view of the overwhelming evidence that the Ratites are secondarily descended from flying birds, the fact that the Ratites already show THE EVOLUTION OF RATITES 67 neoteny in two other features, the plumage and the bones of the skull, and the probability, from A. Kleinschmidt’s (1951) reconstruction, that the palate of Archaeopteryx was neognathous (schizognathous), it is impossible to believe that in their palates the Ratites are primitive. The palaeognathous type of palate must therefore be neotenous. This means a complete reversal of the hitherto generally held view of the palate of birds and necessitates the conclusion that the so-called neognathous palate is primitive. The primitive nature of the neognathous palate in birds is probably connected with the phenomenon of kinetism. J. Versluys (1910) has shown that the mesokin- etic condition in Carinates, where the quadrate and pterygoid bones are capable of a certain amount of movement and sliding, and there is a joint between the pterygoid and palatine whereby the upper jaw can be moved on a hinge at the level of the lacrimal bones and raised relatively to the brain-case, is only intelligible if the birds were evolved from reptiles in which a similar though less extensive power of movement was possible: the condition which he has called metakinetic. According to him (IgI0 : 244) even Archaeopteryx had a kinetic skull capable of movement, although it still possesses a preorbital bar separating the preorbital fossa from the orbit, and a suborbital bar. But in the Ratites the power of movement has been reduced ; the quadrate has a broad connexion with the pterygoid, the latter has equally broad connections with the palatines and the prevomer, and there is no movable joint between the pterygoid and the hemipterygoid because these two elements have not become separated. It must be concluded therefore that with the loss of flight, general increase in size, and acquisition of different feeding habits, the Ratites have lost the Carinates’ power of movement of the upper jaw, by retaining the juvenile condition of the palate before any joint is formed. I am greatly indebted to Dr. W. C. Osman Hill for informing me that even in the kiwi, which is the smallest of the Ratites, there is no mobile joint at the base of the upper jaw; and that in the cassowary the only very slight mobility in the upper jaw is at a point far forward, just behind the nostrils. Further, there is a curious point in the distribution of the palaeognathous type of palate among the birds. It is found not only in Ratites, but also in the tinamus, which are Carinates with a well-developed keel on the sternum and good power of flight. This fact in itself is sufficient to indicate that the Ratites have lost the power of flight, for it could hardly be contended that the tinamus have evolved flight from a flightless Ratite condition. As for the argument that the absence of any rudiment of the clavicle in the ostrich implies that it was evolved from ancestors which lacked the clavicle (and, by implic- ation, could not fly), it is another example of the fallacies to which the theory of recapitulation leads by its assertion that early embryonic stages of development must represent early ancestral stages in evolution. Modern birds lack even the rudiments of teeth, but teeth are present in Archaeopteryx, Hesperornis, and Ichthyornis. The absence of tooth rudiments in modern birds no more excludes Archaeopteryx from their ancestry than the absence of limb-rudiments in snakes indicates that their ancestors were limbless. Finally, the embryonic development of the emu, studied by H. Steiner (1936) 68 THE EVOLUTION OF -RATITES and H. Lutz (1942), shows that the structure and organization of the Ratite embryo is so similar to the Carinate that it can only be interpreted on the view that Ratites have evolved from flying birds. CONCLUSIONS On all these grounds, therefore, there can be no doubt that Owen was correct in regarding the Ratites as birds which have “ abrogated’ the power of flight. It is possible to go further and to say that they have degenerated from a Carinate condition. Whether the Ratites represent a natural group or whether they are an assemblage of forms which have independently followed parallel lines of evolution consequent on the loss of flight is a further problem for ornithologists to solve. In view of the incontrovertible evidence from the structure of the wing, the pygostyle, and the cerebellum, that the Ratites have degenerated from flying birds, any attempt to explain the persistent juvenile characters of the Ratites (nestling- down, skull-sutures, and palate) as phylogenetically primitive is doomed to failure ; and the Ratites must be regarded as providing one of the most telling exposures of the fallacy of the theory of recapitulation. I am glad to acknowledge the help of my colleagues in the Bird Room of the British Museum (Natural History), Mr. J. D. Macdonald and Miss P. A. Lawford, of Dr. W. E. Swinton of the Department of Geology, and of Mr. J. V. Brown, Senior Photographer. SUMMARY Now that the anatomy of Archaeopteryx is adequately known, it is possible to make a rigorous analysis of the characters of the Ratites in the light of the conditions shown by primitive birds. The structure of the wing, tail, and brain in Carinates shows advances on Archaeopteryx which are undoubtedly adaptations to flight. The presence of the same features in Ratites proves that they are descended from flying birds. The condition of the plumage, the sutures between the bones of the skull, and the disposition of the bones of the palate in Ratites, all show secondary retention of characters which are juvenile in Carinates, and are evidence of neoteny in the Ratites. REFERENCES DE BEER, G. R. 19544. Archaeopteryx and Evolution. The Advancement of Science. London, 42 : 160-170. 1954b. Archaeopteryx lithographica. London, i-xi, 64 pp. Broom, R. 1906, On the early development of the appendicular skeleton of the Ostrich with remarks on the origin of birds. Tvans. S. Afr. Phil. Soc., Cape Town, 16 : 355-368. 1947. Did the ancestors of the ostrich fly? Ostrich, Pretoria, 18 : 47-50. CHANDLER, A.C. 1916. A study of the structure of feathers with reference to their taxonomic significance. Berkeley Univ. Cal. Pub. Zool. 13, No. 11 : 243-446. Darwin, C. 1859. Origin of Species. London. DuERDEN, J. E. 1920. Method of the degeneration in the Ostrich. Journ. Genetics, Cam- bridge, 9 : 131-193. EpIncER, L. 1912. Uber das Kleinhirn und den Statotonus, Dtsch. Z. Nervenheilk., Leipzig, 45 : 300-311, THE EVOLUTION OF RATITES 69 EDINGER, T. 1941. The brain of Pterodactylus. Amer. Journ. Sci., New Haven, 239 : 665- 682. Ewart, J.C. 1921. The nestling-feathers of the Mallard, with observations on the composi- tion, origin and history of feathers. Proc. Zool. Soc. London, 1921 : 609-642. FrRIANT, M. 1945. Les carpiens du Nandou (Rhea). C. R. Acad. Sci. Paris, 221 : 641-643. 1945. Developpement et interprétation de la ceinture scapulaire du Nandou (Rhea). C. R. Acad. Sci. Paris, 221 : 711-713. 1946. Le Procoracoide des Oiseaux. C. R. Acad. Sci., Paris, 222 : 153-155. FURBRINGER, M. 1888. Untersuchungen zuy Morphologie und Systematik dey Vogel. Amster- dam, 2 vols. xlix, 1751 pp. Grecory, W. K. 1935. Remarks on the origins of the Ratites and Penguins. With discus- sion by Robert Cushman Murphy. Proc. Linn. Soc. New York, Nos. 45-46 : 1-18. Hux.ey, T. H. 1867. On the classification of birds and the taxonomic value of the modi- fication of certain of the cranial bones observable in that class. Pyoc. Zool. Soc. London : 415-472. Inevar, S. 1918. Zur Phylo-und Ontogenese des Kleinhirns. Folia Neuro-Biologica, Haarlem, 11 : 205-495. Kiernscumipt, A. 1951. Uber eine Rekonstruktion des Schadels von Archaeornis siemensi Dames 1884 im Naturhist. Mus. Braunschweig, Int. Ovn. Congr. 10, 1950 (1951) : 631- 635- Linpsay, B. 1885. On the avian sternum. Pyvoc. Zool. Soc. London : 684-716. Lowe, P. R. 1928. Studies and observations bearing on the phylogeny of the Ostrich and its allies. Pvoc. Zool. Soc. London : 185-247. 1935. On the relationship of the Struthiones to the Dinosaurs and to the rest of the avian class, with special reference to the position of Aychaeopteryx. Ibis, London, (13) 5: 398-432. 1944. An analysis of the characters of Archaeopteryx and Archaeornis. Were they reptiles or birds? Jbis, 86: 517-543. Lutz, H. 1942. Beitrag zur Stammesgeschichte der Ratiten. Vergleich zwischen Emu- Embryonen und entsprechendem Carinatenstadium. Rev. suisse Zool. Genéve, 49 : 299- 399. MarsHatt, W. 1873. Beobachtungen iiber den Vogelschwanz. WNiedevl. Arch. Zool., Haarlem & Leipzig, 1 : 194-210. MAYNARD SMITH, J. 1952. The importance of the nervous system in the evolution of animal flight. Evolution, Lancaster, Pa., 6: 127-129. McDowELl, S. 1948. The bony palate of birds. Part I: The Palaeognathae. The Auk, 65 : pp. 520-549. OweEN, R. 1866. On the Anatomy of Vertebrates. 11. Birds and Mammals. London, i-viii, 586 pp. ParRKER, T. J. 1892. Observations on the anatomy and development of Apieryx. Philos. Tvans. B. 182, 25-134. Parker, W. K. 1875. On the morphojogy of the skull in the Woodpeckers (Picidae) and Wrynecks (Yungidae). Tyvans. Linn. Soc. London, Zool., 1: 1-22. 1876. On the structure and development of the bird’s skull. Tvans. Linn. Soc. London, Zool., 1: 99-154. 1877. On Aegithognathous birds (Part I). Tyvans. Zool. Soc. London, 9 : 289-352. 1879. On the skull of Aegithognathous birds (Part II). Tyvans. Zool. Soc. London, 10 : 251-314. Pycrart, W. P. 1900, On the morphology and phylogeny of the Palaeognathae (Ratitae and Crypturi) and Neognathae (Carinatae). Tvans. Zool. Soc. London, 15 : 149-290. 1900. On the palate of Caprimulgidae. Bull. Brit. orn. Cl. xi: 12-13. 1go1. Some of the points in the morphology of the palate of the Neognathae. Journ. Linn. Soc. London, Zool., 28 : 343-357. Smupson,G.G. 1946. Fossil Penguins. Bull. Amey. Mus. nat. Hist. 87 : 7-99. 70 THE EVOLUTION OF RATITES STEINER, H. 1918. Das Problem der Diastataxie des Vogelfliigels. Jena Z. Naturw.: 55 : 221-246. 1936. Ueber die aiissere Gestaltung eines fiinfzehntagigen Embryos des Emus. Rev. suisse Zool. Genéve, 43 : 543-550. 1938. Der ‘ Archaeopteryx’’ Schwanz der Vogelembryonen. Vjschy. naturf. Ges. Ziivich, 83, Beiblatt : 279-300. 1949. Zur Frage der ehemaligen Flugfahigkeit der Ratiten. Rev. suisse Zool. Genéve, 56 : 364-370. — 1956. Die taxonomische und phylogenetische Bedeutung der Diastataxie des Vogel- fliigels. J. Ovn., Lpz., 97: 1-20. STRESEMANN, E. 1927-1934. Sauropsida; Aves, etc., xi, 899 pp. in Kuekenthal, Handbuch der Zoologie, 7 : Halfte II. TuckER, B. W. 1938a. Functional evolutionary morphology: the origin of birds. Evolu- tion, Essays on Aspects of Evolutionary Biology : edited by G. R. de Beer, Oxford, 321-336. 1938). Some observations on Dr. Lowe’s theory of the relationship of the Struthiones to the Dinosaurs and to other birds. Proc. 8th Int. orn. Congr. Oxford, 1934 : 222-224. VERSLUYS, J. 1910. Streptostylie bei Dinosaurien, nebst Bemerkungen itiber die Verwand- schaft der Vogel und Dinosaurier. Zool. Jahyb. Jena (Abt. Anat. & Ontogenie d. Tiere) : 30 : 175-260. Wray, R. S. On some points in the Morphology of the Wings of Birds. Pyoc. Zool. Soc. London : 343-357- Younea, J. Z. 1950. The Life of Vertebrates. Oxford. EXPLANATION OF PLATES PLATE 5 (1) Right side view of the pygostyle in a Carinate, Leptoptilos cywmeniferus (Marabou Stork) and (2), ina Ratite, Stvuthio camelus, showing the similarity of structure. (ap), the anterior portion showing the elements ofa distinct vertebra ; (pp) posterior portion composed of fused vertebrae. PLATE 6 (1) The brain as seen in right-side view in Archaeopteryx lithogvaphica. (2) sagittal section of the brain in a Carinate, Tvinga ocrophus (Green Sandpiper), and (3) in a Ratitie, Rhea americana. (ce), cerebellum ; (ch), cerebral hemisphere ; (ol), optic lobes. PLATE 7 The arrangement of the feathers on the wing of a young Ratite, Rhea americana, showing the differen- tiation between primary and secondary remiges, bastard wing, and wing-coverts. PLATE 8 (1) Ventral view of the structure of the palate in the Carinate Pygocelis papus (Gentoo penguin) nestling ; (2) in the Carinate Anthvopoides parvadisea (blue crane) ; and (3) in the Ratite Dromiceus novae-hollandiae (emu). (hpt), hemipterygoid ; (pa), palatine ; (pt) pterygoid; (pr), prevomer; (qu), quadrate. PLATE 9 (1) The structure of the palate as seen in left-side view in Corvus frugilegus (rook) young ; (2), Mega- laema virens (Himalayan barbet). (hpt), hemipterygoid ; (ju), jugal; (pa), palatine; (pt), pterygoid ; (pr), prevomer ; (qu), quadrate. Bull, B.M. (N.H.) Zool. 4, 2 PLATE 5 PLATE 5 (t) Right side view of the pygostyle in a Carinate, Leptoptilos crumeniferus (Marabou Stork), and (2), in a Ratite, Struthio camelus, showing the similarity of structure. (ap), the anterior portion showing the elements of a distinct vertebra ; (pp) posterior portion composed of fused vertebrae. Bull. BM. (N.H.) Zool. 4, 2 PLATE 6 PATE 6 (1) The brain as seen in right-side view in Archaeopteryx lithographica. (2) sagittal section of the brain in a Carinate, Tvinga ocrvophus (Green Sandpiper), and (3) ina Ratite, Rhea ameri- cana. (ce), cerebellum ; (ch), cerebral hemisphere ; (ol), optic lobes. HA ke kK ¢ P >, (* > » L te toad 4 5 \ a N Bull. B.M. (N.H.) Zool. 4, 2 PEATE 7 ring the vyicana, Sho and wing-coverts. “ w c a = = oa 20 es = oy ~~ n we R Sate) nr V a0 Sp mS oS pa <q Secondaries differentiation between primary and secondary remi The arrangement of the feathers on the w Bull. B.M. (N.H.) Zool. 4, 2 PLATE 8 PLATE 8 (1) Ventral view of the structure of the palate in the Carinate Pygocels papus (Gentoo pen- guin) nestling ; (2) in the Carinate Anthvopoides paradisea (blue crane) ; and (3) in the Ratite Dromiceus novae-hollandiae (emu). (hpt), hemipterygoid; (pa), palatine; (pt) pterygoid ; (pv), prevomer ; (qu), quadrate = Bull. B.M. (N.H.} Zool. 4, 2 PLATE ju pa pt Zi PLATE 9 (1) The structure of the palate as seen in left-side view in Corvus frugilegus (rook) young; (2), Megalaema virens (Himalayan barbet). (hpt), hemipterygoid ; (ju), jugal ; (pa), palatine ; (pt), pterygoid ; (pr), prevomer ; (qu), quadrate. 9 PRINTED IN GREAT BRITAIN BY ADLARD AND SON, LIMITED, BARTHOLOMEW PRESS, DORKING DENYS W. TUCKER . ? ene Wega. ‘ é BULLETIN OF BRITISH MUSEUM (NATURAL HISTORY) Vol. 4 No. 3 ~LONDON: 1956 STUDIES ON THE TRICHIUROID FISHES—3 A PRELIMINARY REVISION OF THE FAMILY TRICHIURIDAE BY DENYS W. TUCKER Pp. 73-130; Pl. 10; 23 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 4 No. 3 LONDON: 1956 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, 1s issued in five series corresponding to the Departments of the Museum, and an Historical Series. Parts appear at irregular intervals as they become veady. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. This paper is Vol. 4, No. 3 of the Zoological series. PRINTED BY ORDER OF THE TRUSTEES OF THE BRITISH MUSEUM Issued August, 1956 Price One Pound STUDIES ON THE TRICHIUROID FISHES—3* A PRELIMINARY REVISION OF THE FAMILY TRICHIURIDAE By DENYS W. TUCKER CONTENTS Page INTRODUCTION . ‘ : e : c = > 5 74: THE CHARACTERS OF THE FAMILY TRICHIURIDAE . : 6 ° a. 7A A Suort Key To THE SUBFAMILIES AND GENERA OF THE Fairy TRICHI- URIDAE . : ‘ é 2 c 4 5 : a Sa SysTEmaTic REVIEW : Subfamily Aphanopodinae : ; a 2 > : : 0 Hl Genus Diplospinus . 4 ° 4 5 5 : 2 7S » Aphanopus . : i i A 5 ‘ ‘ A 81 », Benthodesmus : : : 4 ‘ . » f 85 Subfamily Lepidopodinae 5 6 5 c : = c - 89 Genus Lepidopus . : - C : : : - - go » Evoxymetopon : : A F F 5 o Cy) » Eupleurogrammus . F ° c 2 5 - I02 » Assurger é 6 < . 6 “ - I06 » TLentoriceps . 9 5 é : 4 a : LTO! Subfamily Trichiurinae . : c : : ; : = = BE: Genus Trichiurus . 5 5 c é 5 4 5 + Lrg », Lepturacanthus : 5 ¢ “ . : a = LO: THE OriciIn, EvoLution AND CLASSIFICATION OF THE TRICHIURIDAE : Summary of earlier work f : 0 6 A 5 . 5 He) Nesiarchus-Diplospinus : the Gempylid-Trichiurid bridge . 0 eel 2 Evolutionary trends in the Trichiuridae : é © - 125 Classification of the Trichiuridae 4 é 3 5 9 5 neh) REFERENCES 5 F ; ; p 3 Fi 5 : . 5 £20 SYNOPSIS This paper presents a working classification of the Trichiuridae, based on a consideration of the literature of the family and examination of selected material, which has been prepared as a prelude to Reports on the “ Dana”’ collections of Trichiuridae and Gempylidae. Three subfamilies are recognized : Aphanopodinae (genera Diplospinus, Aphanopus, Benthodesmus) ; radiation of the Trichiuridae are discussed. 1 The previous papers in this series were: (1) The fishes of the genus Benthodesmus (Family Tri- chiuridae), Proc. zool. Soc. Lond. (123 : 171-197, 3 pls., 5 text-figs. (1953). (2) Benthodesmus tenuis ZOOL, 4, 3. 6 74 THE FAMILY TRICHIURIDAE INTRODUCTION As I contemplated the mass of material which resulted from my rash acceptance of Dr. Anton Fr. Bruun’s invitation to write reports on the young Trichiuroid fishes collected by the “‘Dana ’’ Expeditions, I realized the urgent need of some preliminary working classification with which to regulate the chaos that must ensue once these many thousands of specimens were released, like so many djinns, from their tubes and bottles. The problem of the Gempylidae was immediately relieved by Matsubara and Iwai (1952) and by Mrs. Marion Grey (1953), but the case of the Trichiuridae remained desperate. There has been no comprehensive revision of this family since the end of the nineteenth century. The earlier synopses of Giinther (1860), Gill (1863) and Goode and Bean (1895) are no longer adequate accounts even of the genera which they describe and, moreover, contain no attempt at a phyletic classification since they date from a period before the planting of family trees became fashionable. Later workers have had varying success in distinguishing the genera and species of limited regions. In this century a few new species and genera have been proposed, two of the latter without any of the inhibitions consequent upon an interest in the family or the possession of study-material. The present draft revision assigns a place to every nominal genus and species and gives, as a minimum, the reference for the first publication of every name and name-combination, together with selected items from the remaining literature. It gives diagnoses and a phyletic classification of all sub-families, genera and species recognized and argues the case for synonymies with whatever detail the individual circumstances may immediately demand. Except for Evoxymetopon, Assurger and Tentoriceps (of which material or new published descriptions would be greatly appreciated), material of all genera and species has been examined, including a substantial number of type specimens. The author of any “ preliminary ”’ contribution should justify his title. The amount of labour involved in preparing the present MS as a working tool has shown the need of such a tool and of certain small but critical contributions to the understanding of the Trichiuridae which those possessing rarer material may make. It will be some considerable time before the final ““Dana ’’ Reports on the Trichiuridae and Gempy- lidae can be completed and so, faute-de-mieux, a preliminary account appears likely to be useful, even though some of its conclusions may be subject to second thoughts. I wish to express my thanks to Messrs. P. E. Purves and A. C. Wheeler of the British Museum (Natural History) for numerous radiographs which have been of very great assistance in this work. THE CHARACTERS OF THE FAMILY TRICHIURIDAE Regan (1909) allies the Trichiuridae with the Gempylidae as the Trichiuriformes, forming the first division of his suborder Scombroidei of the order Percomorphi. He characterises the Trichiuriformes as having :— “Caudal fin-rays not deeply forked at the base, the hypural in great part THE FAMILY TRICHIURIDAE 75 exposed. Praemaxillaries beak-like, free from the nasals; mouth toothed, with lateral cleft; strong anterior canines. Epiotics separated by supra- occipital. Gill-membranes free from the isthmus. Pectoral fins placed low.” With this diagnosis I have no present disagreement save to comment that hypurals are sometimes absent and to prefer the use of “‘fangs’”’ or ‘‘ caniniform teeth ” rather than ‘‘canines’’ for fish teeth; the term ‘‘canine’’ is best restricted to certain reptiles and to the mammals, in which it is defined, not by form but by position and homology, as ‘‘ the most anterior tooth of the maxilla, situated on or immediately behind the premaxillo-maxillary suture . . . or the tooth in the lower jaw which bites in front of the upper canine ”’. ce Regan’s diagnosis of the family Trichiuridae follows : “Body very elongate, strongly compressed; maxillary sheathed by the praeorbital ; spinous dorsal, if distinct, not longer than the soft!; anal with numerous short spines? ; pelvic fins reduced to a pair of scale-like appendages or absent? ; caudal small or absent. Dorsal and anal rays corresponding to the vertebrae‘, each interneural or interhaemal attached to a neural or haemal spine ; pelvic bones, if present, united to form a slender spicular bone connected with the cleithra by a long ligament®. Vertebrae numerous, 100(43 + 57) to 159(39 + 120) or more®; ribs feeble, sessile.” This description is evidently based primarily upon examinations of Lepidopus, Aphanopus and Trichwurus and requires several modifications and qualifications : (rt) The spinous dorsal is always distinct ; it is longer than the soft in Diflospinus (discovered since Regan’s time) and very slightly longer than the soft in occasional specimens of Aphanopus. (2) Some, if not all, of the anal rays are split, soft and support a fin-membrane (Diplospinus, Aphanopus, Benthodesmus, Lepidopus, Evoxymetopon, Assurger) ; in Trichiurus, Lepturacanthus and Eupleurogrammus, however, the anal rays are much reduced spinules or entirely absent. At the origin of the anal fin, moreover, immediately behind the vent, are two spines (represented by the notation i+ I throughout the present paper) ; of these the anterior is a minute spinule while the second may be variously enlarged as a leaf-like or keeled scute, or as a stout spine. (3) The pelvic fins in some genera (Diplospinus, Aphanopus, Benthodesmus, Lepidopus) and probably in all in which they are present, consist each of a scale-like spine and one rudimentary soft ray, the latter newly noticed. (4) The dorsal spines and their basals and interneurals always correspond to the trunk vertebrae ; the dorsal soft rays may be twice as numerous as the adjacent vertebrae (Diplospinus), slightly more numerous (Aphanopus, Benthodesmus) or as numerous (remaining genera). (5) The pelvic bones form an imperfectly fused, fenestrated structure which is not always elongated. (6) The vertebrae range from 34 + 24 = 58 (Diplospinus) to 53 + 103 = 156 (Benthodesmus simonyt) or 41 + 151 = 192 (Eupleurogrammus muticus). THE FAMILY TRICHIURIDAE 76 “QABOUOD SI IE[N -rodoqns oy} JO UlsIeUI 9a1f OL -odeu oy} 0} peuy -uoo st ‘Aprumeyqns sty} Ur “Yor “4seI9 eyes & WIIOF 0} payeAaya SI SeSprr [e}UOIF ot]} JO GoUEN|IOS iot1eysod ayy, *(snaniyotaz) pesy ULINTyOU yeordAy y—'? ‘SI *xaAuo09 st repnoredoqns ay} JO UISIeUL 9e1F OY L -adeu oY} 0} pauyuos ‘IaABMOY “WLIO SI} UT ‘ST YOTYA “4seI9 yez18es © UIOJ OF Payer -a]9 SI SoSplI [e}WoIF 9Y} JO eoueNBUos rotsaysod ay ‘sndopidaT : peey autpodopidey jo sadA,—'z “Oly “x9Auod st IepNoredoqns ay} Jo espa very OUL “uoIse1 [ewUOF-owyje orUe oy} Jo uoHeAeTa UP se jNOUS ayy Suope preMsoy panuyjUoo st YOST ‘S919 ye} 18es ® WLOF 0} poyeAgfa St sasprt [e}UOIF oy} FO souON -uo9 ro1eysod ayy, “(weag 2 ePOOH Ie7je uMeIp-d1) uodojamAxong : peay eutpodoprideT yo sodA, —'€ ‘91 *x9Auo0o st 1e[nosedoqns 94} JO UISTeU aerF OY} £4so10 © ULIOF jou saop sespit [e}UOIZ VY} FO aoUENYUOS Iotieysod ay} { UOrsar [e}UOIF-OUIYZE OY} FO UOTPeAQIE OU ST 99 L ‘(sndounydp) peay autpodoueydy pestdAy y—TI “ST THE FAMILY TRICHIURIDAE 77 A SHORT KEY TO THE SUBFAMILIES AND GENERA OF THE FAMILY TRICHIURIDAE Frontal ridges not elevated, no sagittal crest. Profile of head rising very gently from snout tip to dorsal (cf. Text-fig. 1) : . | APHANOPODINAE (Pp. 77) D.72-73. Spinous dorsal base twice as long as s soft. : 2 . Diplospinus (p. 78) D.82-87. See “ Lepidopus xantusi ’’ (Lepidopodinae) D.g1-95. Spinous and soft dorsal bases sub-equal . : : . Aphanopus (p. 81) D.120+ Spinous dorsal base half as long as soft . c¢ 5 . Benthodesmus (p. 85) Posterior confluence of frontal ridges elevated, forming a prominent sagittal crest at the nape, which may or may not be continued forward as a ridge-like elevation of the ethmo-frontal region (cf. Text-figs. 2, 3 and 4). Ventral fins present. Lateral line descending gently from the shoulder and median or sub-median along the body, i.e. distance from lateral line to ventral profile at anus much more than half distance from lateral line to dorsal. Lower hind margin of operculum convex . 2 c c 4 LEPIDOPODINAE (p. 89) Sagittal crest confined to nape. Interorbital concave. Caudal present Lepidopus (p. 90) Sagittal crest continuous from snout tip to dorsal. Interorbital convex Caudal present D.87-93. Body depth 12-13 in length . : C : . Evoxymetopon (p. 97) D.120. Body-depth 20-28 in length . g z c a . Assurger (p. 106) Caudal absent. Body depth 14-18 in length F ; 7 2 : Eupleurogrammus (p. 102) Body depth 20-24 in length : é Tentoriceps (p. 110) Ventral fins absent. Lateral line descending steeply from the shoulder and running near the ventral profile of the body, i.e. distance from lateral line to ventral profile at anus less than half distance from lateral line to dorsal. Lower hind margin of operculum more or less concave. Caudal always absent (cf. Text-fig. 4) TRICHIURINAE (p. 112) Post-anal scute small, less than the pupil. Soft anal rays not breaking through skin. Eye large, 5.0-7.0 in head 3 5 c Trichiurus (p. 113) Post-anal scute large, half the eye-diameter. ‘Soft ‘anal Tays pungent spinules, breaking ventral profile. Eye small, 6.7—10.0 in head . . Lepturacanthus (p. 119) SYSTEMATIC REVIEW Subfamily APHANOPODINAE Gill Aphanopodinae Gill, 1863, Proc. Acad. nat. Sci. Philad. 1863 : 225. Type genus Aphanopus Lowe. GENERA NOW RECOGNISED.—Aphanopus Lowe ; Benthodesmus Goode & Bean ; Diplospinus Maul. Dracnosis : A. Snout gently sloping ; orbits entering upper profile of head ; frontal ridges only slightly elevated, not contributing to a sagittal crest. B. A stout, conical, cartilaginous protuberance at the mandibular symphysis ; another, much smaller, at the tip of the snout. C. Lower hind margin of operculum markedly convex, 78 THE FAMILY TRICHIURIDAE D. Teeth of main series with double barbs (Diplospinus) or entirely without barbs (Aphanopus, Benthodesmus). E. Teeth on palatines in a linear series. (In Aphanopus only 1-2 posterior rudiments of the series present.) F. Lateral line descending gently from the shoulder and running in a median or sub-median position along the body, i.e. distance between lateral line and ventral profile much more than half distance between lateral line and dorsal. G. Spinous dorsal fin long, with 32-46 rays. Spinous and soft dorsals partly divided by a slight notch. H. Soft dorsal rays slightly more numerous than adjacent caudal vertebrae, or up to twice as many. Basal and interneural elements intercalated among the main series and unrelated to neural spines of vertebrae. I. Spinous anal i+ 1; anterior soft anal rays weak but (except in Benthodesmus simonyt) an external fin is continuous in some form or other from the vent nearly to the caudal; the properly developed fin may extend the whole length or be confined to the posterior 20-25 rays. Terminations of dorsal and anal fins sub-opposite. . Caudal fin always present ; small, normal, forked. Ventral fins always present (though reduced to internal rudiments in adult Aphanofus), composed each of a scale-like spine and one soft ray ; in the adult fish inserted not more than 2-3 mm. before/behind anterior/posterior perpendiculars through the ends of the pectoral base. M. Pyloric caeca few (6-9) ; (not verified in Diplospinus). en Osteological Literature Giinther, 1860, Cat. Fish. B.M. 2 : 342-344 (desc. osteology Aphanopus). Tucker, 1953, Proc. zool. Soc. Lond. 123 : 196-197, pls. 2-3 (figs. osteology of paired fins and anal fin of Aphanopus & Benthodesmus). 1955, Bull. Mus. Hist. nat. Belg 31, No, 64: 1-26 (figs. osteology of pelvic and anal fin of Benthodesmus). Literature on young stages Maul, 1948, Bol. Mus. Funchal No. 3, Art 6 : 42, fig. 17 (young Diplospinus). Tucker, 1953, op. cit. : 187 (figs. young Aphanopus and Benthodesmus). Genus DIPLOSPINUS Maul Diplospinus Maul, 1948, Bol. Mus. Funchal No. 3, Art. 6: 42. Type species Diplospinus multistriatus Maul. Monotypic. Synonyms Lepidopus (non Gouan 1770) (part) Brauer, 1906. Benthodesmus (non Goode & Bean 1882) (part) Goode & Bean, 1895 ; Fowler, 1938. (Refs. below.) THE FAMILY TRICHIURIDAE 79 Diagnosis : (t) Body elongate, head length 6-6-6-9 in standard length 125-203 mm., body depth 18-5 in S.L. (2) Vent exactly in middle of S.L. (3) Vertebrae 34 + 24 = 58. (Corresponds to 36 + 22 = 58 in convention used for Benthodesmus.) (4) Spinous dorsal base twice as long as soft dorsal base. (5) Dorsal spines 32-33 ; dorsal soft rays 40. 6) Dorsal soft rays about twice as numerous as adjacent caudal vertebrae, so that alternate interneural elements do not articulate with neural spines. (7) Anal spines i + I, the former half the length of the latter in young stages ; condition in the adult unknown; iis linear ; I is dagger-shaped and V-shaped in transverse section. (8) Anal spines i and I articulate close together on a common basal, which is not enlarged or specially modified and which, except that it does not quite touch the corresponding haemal arch, does not show any difference in the size and relations of the interhaemal process from those which follow it. (Condition similar to Lefidopus.) (9) A complete external anal fin supported by 31 split but unbranched rays extending from the spinous anal nearly to the caudal. The soft rays and their basal elements are about twice as numerous as the adjacent caudal vertebrae, so that alternate basals have interhaemal processes which are unrelated to haemal arches. (10) Ventral fins inserted on perpendicular through anterior end of pectoral fin-base. (rz) Ventral fin I-r ; a narrow scale-like spine and an external split ray twice as long. (12) All principal teeth of the premaxillary and dentary series are strongly barbed (arrowhead-shaped), with thickened enamel caps. (13) Palatine teeth in a linear series, exposed. (14) Principal teeth on first gill-arch numerous. (15) Long intermuscular (pleurals and epipleurals) bones present, extending throughout trunk. (16) Melanophores distributed in parallel and narrow longitudinal rows along the body. One species, Diplospinus multistriatus Maul, Atlantic and Indo-Pacific. Diplospinus multistriatus Maul, (Text-fig. 5) Benthodesmus atlanticus (part) Goode & Bean, 1895, Oceanic Ichthyology : 206 (the two small specimens mentioned, fide Dr. Carl L. Hubbs, in Uitt.). Benthodesmus benjamini (part) Fowler, 1938, Proc. U.S. Nat. Mus. 85 : 45 (certain of the para- types, fide Dr. Carl L. Hubbs, in litt.). ? Lepidopus gracilis Brauer, 1906, Wiss. Evgeb. “‘ Valdivia” 15 : 291, Taf. XII, fig. 1 (not fig. 5 as erroneously stated in the text nor fig. 3 as stated in the legend to the plate). Holotype in the Berlin Museum? Type locality West coast of S, Africa, St. 82, 21° 53’ S., 6° 58’ 6” E, THE FAMILY TRICHIURIDAE so (pezeqye ‘ gh6r ‘[nepy 197F¢V) “TS ‘wu €oz ‘adAjojoHH [NeW SNH2AQS1q]NUL snurdsojgiq—S “oI *HD OF THE FAMILY TRICHIURIDAE 81 Diplospinus multistriatus Maul, 1948, Bol. Mus. Funchal, No. 3, Art 6: 42, fig. 17. Holotype Museu Municipal do Funchal No. 3063. Type locality Madeira. Paratypes Museu Municipal do Funchal Nos. 3064-5, 3067-9. Paratype British Museum (Natural History) No. 1953.10.28.1. (Formerly 3066.) Certain discrepancies will be noticed between the generic diagnosis given above and the otherwise accurate description and figure by Maul (1948) ; the corrected observations have been made on the paratype kindly presented by Mr. G. E. Maul. Each ventral fin includes a soft ray in addition to the spine ; there is a single row of about a dozen teeth on each palatine (“‘ no teeth on vomer or palatines’”’) ; there are traces of an apparent and highly probable lateral line (“ no lateral line ’’) though the present specimen is completely skinned ; certain of the premaxillary fangs are represented by replacement teeth (“ depressible teeth ’’). The number of branchio- stegal rays is 7, as in other Trichiurids. The number of pyloric caeca cannot be determined owing to destruction of the thoracic region. There is a deep notch on the hinder margin of the opercular, as already observed by Maul, and this character proves to be rather important since it is confined to Diplospinus, the most primitive recent Trichiurid and to Neszarchus, the nearest-related Gempylid (see p. 124). Since Brauer’s (1906) figure of Lepidopus gracilis bears the magnification 2/1 we may deduce a S.L. of 68 mm., i.e. about one-third the length of the type series of Diplospinus multistriatus. The head is 4:8 and the height 14-4 in the length; the eye goes 5 times in the head, and the ventral and anal spines are proportionately longer than in the types. All these differences are in the directions to be expected in a younger fish. The counts of D.65-67 and A.27 are slightly low, but not outside the probable range of variation or error. However, the eye is shown about a quarter its diameter below the dorsal profile of the head, the origin of the dorsal fin is a little retarded and the insertion of the ventral fins is below the posterior rather than the anterior end of the pectoral base (“ Bauchflosse kurz hinter der Vertikale der Brustflosse ’’). These discrepancies must await a satisfactory explanation, which is likely to result in Diplospinus gracilis (Brauer) becoming the definitive name of the present species. Genus APHANOPUS Lowe Aphanopus Lowe, 1839, Proc. zool. Soc. Lond. 7: 79. Type species Aphanopus carbo Lowe. Monotypic. Synonyms Lepidopus (non Gouan, 1770) Sim, 1898 ;Dons, 1921. (Refs. below). DIAGNOSIS : (1) Body elongate, head length 5-68-4-92 in standard length 102-1036 mm., body depth 21-7-11-23 in same. 2) Tail 48-49% of standard length. 3) Vertebrae 42-44 + 55-56 = 98-99. 4) Spinous and soft dorsal bases sub-equal, differing by at most + 3 % of S.L. 5) Dorsal spines 38-41 ; dorsal soft rays 53-56; aggregate 91-95. 6) Dorsal soft rays practically corresponding with adjacent caudal vertebrae. ( ( ( ( ( THE FAMILY TRICHIURIDAE 82 (ueuntoads oy} w01; paiezye ‘(ghO1) [ney Joye ‘Tey snow py jo adAjojoH ) “T'S ‘wu zor apuaant ‘amoT ogtva sndounydyp—L “diy WIC “T'S ‘wur zio‘r ‘adAjopoyy «amo 0gav9 sndouvydp—'9 “91g THE FAMILY TRICHIURIDAE 83 There are very few intercalated interneural elements which are unrelated to neural spines and these occur usually towards the beginning and end of the fin. (7) Anal spines i + I, the former about 1/5 the length of the latter in the young stages but becoming disproportionately smaller in the adult, in which i becomes a minute sharp spinule, usually concealed beneath the skin and I is a stout dagger-shaped spine, triangular in cross-section. (8) Anal spine i articulates a short distance in advance of I. Their common basal element isa complex, greatly enlarged and strengthened to accommo- date the hypertrophied I, and representing four or more fused elements. The compound interhaemal process is stout and does not touch the adjacent haemal arch. The horizontally directed component of the compound basal element occupies the length of three vertebral centra and the presumed anterior migration of the corresponding interhaemal processes leaves a space above it. (9) A complete external anal fin of 44-48 split but unbranched rays extending from the spinous anal nearly to the caudal. The anterior rays are very weak and the functional fin is effectively confined to the posterior 20-25 rays. The internal supporting skeleton is quite regular ; there is a precise correspondence between rays, basal elements and caudal vertebrae, with a close association between interhaemal processes and haemal arches. (10) Ventral fins inserted immediately before perpendicular through anterior end of pectoral fin-base. External fins present only in the juvenile ; fins and girdle reduced to an internal rudiment in the adult. (rz) Ventral I-r in the juvenile only ; a narrow spine and an external split ray initially about 3 times as long. (12) Principal teeth of the premaxillary and dentary series without barbs: if these are sometimes present on the premaxillary fangs they are usually barely perceptible and confined to the hinder edges, without enamel thickening. The marginal teeth of the jaws are stout, triangular and have microscopically-serrated edges. (13) Palatine teeth reduced to I-2 minute rudiments at hinder end of bone, very much concealed. (14) Principal teeth on first gill-arch very numerous. (15) Intermuscular bones (pleurals and epipleurals) weaker than in Diplospinus. (16) Pigmentation uniform, dense ; fish uniform black when dead. Living fish coppery with iridescent reflexions. One species, Aphanopus carbo Lowe, N. Atlantic and Gulf of Aden. Aphanopus carbo Lowe (Text-figs. 6 & 7). Aphanopus carbo Lowe, 1839, Proc. zool. Soc. Lond. 7: 79. Holotype B.M. (N.H.) No. 1851.11.29.6. Type locality Madeira. Aphanopus minor Collett, 1886, Chr. Vid.-Selsk. Forh. 1886 No. 19 : 1, fig. r. Holotype in Universitetets Zoologiske Museum, Oslo. Type locality Denmark Strait, E. of Greenland, 65° N., 31° W. 84 THE FAMILY TRICHIURIDAE Lepidopus caudatus (non Euphrasen, 1788) Sim, 1898, Ann. Scot. nat. Hist. 1898 : 53. Aphanopus schmidti Saemundsson, 1907, Vid. Medd. naturh. Foren. Kbh. 59: 22, Vet ae Holotype in Nattirugripasafnid, Reykjavik. Paratype B.M. (N.H.) No. 1925.7.23-4. Type locality Vestmann Is., S.W. of Iceland. Lepidopus atlanticus (non Goode & Bean, 1895) Dons, 1921, Tvomso Mus. Aarsh. 43, No. 6: 10, fig. I. (Identification corrected to Aphanopus schmidti by Soot-Ryen, 1936, Nytt. Mag. Naturv. 76 : 237.) Aphanopus microphthalmus Norman, 1939, Sct. Rep. John Murray Exped. 7 No. 1: 71, fig. 25. Holotype B.M. (N.H.) No. 1939.5.24.1322. Type locality Gulf of Aden. Aphanopus acus Maul, 1948, Bol. Mus. Funchal No. 3, Art. 6: 47, fig. 18. Holotype in Museu Municipal do Funchal. Type locality Madeira. (Withdrawn as young A. carbo by Maul, 1949, Bol. Mus. Funchal, No. 4, Art. 10: 21.) non Aphanopus simonyi Steindachner, 1891. (See under Benthodesmus simonyi.) non Aphanopus carbo Norman, 1937. (Mediterranean records based on confusion with Lepidopus caudatus, q.v-) TABLE I. Holotype. Vertebrae. Dorsal. Anal. A.carbo . 4 ; 42 +56 5 XXXVIII, 56. 1+1+48 A. schmidti : t 42 +56 6 XXXVIII,55 . i+I+46 A. minor .« : 4 44+? IEA : i+I-+? A. microphthalmus . 44+55 5 XLI, 54 c i+1+45 In the type of A. minor the tail has been broken off a short distance behind the vent and has subsequently healed over with some slight re-orientation of the soft dorsal and anal rays remaining. The remnant includes 25 caudal vertebrae, 28 soft dorsal rays, 21 anal elements. Through the kindness of Dr. C. Stap-Bowitz (Oslo), Dr. Finnur Gudmundsson (Reykjavik) and Mr. G. E. Maul (Funchal) I have been able to examine the types of all the nominal species of Aphanofus and, by comparing these with a series of some thirty specimens from the type locality and as many more from the North Atlantic, to decide that they represent only one species, A. carbo Lowe. A. acus Maul is a juvenile A. carbo and has already been adequately dealt with by Maul (1949). Meristic counts for the other nominal species are given in Table I. Ranges of vertebral counts for the long series are not yet available. but the variations now tabulated are small and well within the limits of those found in Benthodesmus tenuis (p. 88). Fin-ray counts on eighteen Madeiran specimens give ranges ' D.XXXVIII-XL, 53-55 (aggregate 91-95) ; A.i+ 1+ 44-48. The validity of A. schmidti has been much debated, Saemundsson fro, Grieg and others con. The arguments will be dealt with in detail elsewhere ; for the present : it is sufficient to state that the two specimens of A. schmidti show no meristic differences from A. carbo nor any measurable differences in body proportions. The shorter dorsal rays noted by Saemundsson are merely broken; the intangible differences in the contour of the head are due to variations of desiccation and fixation, and may be observed in some of the fishes on the Funchal Market slabs ; the colour described with poetic exactitude by Saemundsson is merely that of a living A. carbo and changes to a glossy black as a post-mortem effect. THE FAMILY TRICHIURIDAE 85 A. minor Collett is founded on a wretched half-grown fish which had somehow contrived to survive the loss of its tail. I have compared the holotype with a Madeiran specimen of equivalent snout-vent length ; there are no differences. A. microphthalmus Norman has been checked against a similar-sized specimen from Madeira ; there are no significant differences. The distension of the branchio- stegal region of the holotype, adequately shown in Norman’s figure. gives an exag- gerated superficial impression of a deeper head and smaller eye. Sim (1898) compares a Scottish fish with Day’s description of Lepidopus caudatus and comments : “Now in the specimen under notice there is not the slightest indication of such ventral scales, and what is considered a scale by the authors named takes the form of a strong, bayonet-shaped spine situated behind the vent, and is an inch long.” Sim clearly had an Aphanopus carbo, at that time unrecognised in the British fauna but since found to be common along the too fathom line, where it may sometimes be taken even by the hundred by vessels trawling for hake. I have a monograph in preparation covering the anatomy and biology of this species. Genus BENTHODESMUS Goode & Bean Benthodesmus Goode & Bean, 1882, Proc. U.S. Nat. Mus. 4: 379. Type species Lepidopus elongatus Clarke. Three species. Goode & Bean erected this genus on the occasion of their describing a fish from Newfound- land which they believed to belong to Clarke’s New Zealand species (the holotype of which they had not seen) and attributed characters to Benthodesmus additional or contrary to those in Clarke’s description. In 1895 (Oceanic Ichthyology : 206) they erected a new species B. atlanticus on their Newfoundland specimen, leaving the situation that Benthodesmus was based on a species which they had not seen. Since the holotype of L. elongatus has been lost I propose to request the International Commission on Zoological Nomenclature to recognize B. atlanticus G. & B. as the type-species of Benthodesmus, which would at the same time provide a more convenient reference point and a more satisfactory indication of Goode & Bean’s intentions. It is practically certain that the two nominal species will eventually be shown to be identical, but for the present I am retaining them both until New Zealand material shall be forthcoming. B. atlanticus is a junior synonym of Aphanopus simonyi Steindachner. Synonyms Lepidopus (non Gouan, 1770) Numerous authors ; for references see under synonymies Aphanopus (non Lowe, 1839) of species. It has been suggested to me that Benthodesmus should be split and a new genus erected on B. tenuis (Giinther). I am strongly opposed to any such action, being of the opinion that B. tenuis is the close ancestor of B. elongatus and that it would be improper to obscure this close relation in the way proposed. In the event of a new genus being recognized there is some possibility of the name Scarcina Rafinesque (1810) being already available, with S. argyrea preceding B. tenuis. Scarcina has always been regarded as a junior synonym of Lepidopus Gouan (1770) and for reasons outlined on p. 94 I prefer to leave it so for the present. THE FAMILY TRICHIURIDAE 86 (‘suoutdads 19430 wod} pejonsysuosar yonur surq) “TS ‘ur 166 ‘adAqo0j077 ‘(19q3UN-D) senuay Snusapoyjuag—6 ‘oI ‘TS wu o/1‘l ‘enopew ‘882° TIT £661 “ON (FN) WE ‘(dauyoepute3s) rdAuours SNuMSapoyjuag—'g “O17 = <A) Tce REET TTT RE SM ee A | THE FAMILY TRICHIURIDAE 87 DIAGNOSIS : (I) Body very elongate ; head-length 7-0-7-6 in standard length 221-591 mm., body-depth 23-8-34:4 in same (B. tenuis) or head-length 6-8-7'8 in S.L. g10-1225 mm., body-depth 21.7-27-0 (B. simony?). (2) Tail 55% (B. tenuis) or 60% (B. stmonyi) of S.L. (3) Vertebrae 47-52 + 75-80 = 123-131 (B. tenuis) or 52-53 + 101-103 = 153-156 (B. simonyt). (4) Spinous dorsal base half as long as soft dorsal base. (5) Dorsal spines 39-42, dorsal soft rays 80-88 (B. tenuis) or dorsal spines 45-46, dorsal soft rays 102-108 (B. simony?). (6) The number of soft dorsal rays is very close to that of the caudal vertebrae. There are very few intercalated interneural elements, which are usually toward the beginning or end of the fin. (7) Anal spines i + I, the former extremely minute and completely concealed in the adult. I is a delicate cardiform scute with a median keel projecting as a short point between the two rounded posterior lobes. (8) Anal spine i articulates a short distance in front of I. Their common basal element is a complex representing three or more fused elements. The interhaemal spine is a thin keel supported by three slender, tubular, cartilage-tipped spines (B. tenuis) or is completely wanting (B. simonyi). The horizontally-directed basal occupies the length of three vertebral centra. (9) A complete external anal fin of 70-76 split but unbranched rays extending from the anal spines nearly to the caudal (B. tenwis) or with the anterior rays wanting and the external fin posterior and reduced to about 25 rays (B. simonyt). (10) Ventral fins inserted immediately before perpendicular through anterior end of pectoral base (B. tenuis) or immediately behind perpendicular through posterior end of pectoral base (B. simonyi). (tz) Ventral fin I, 1 (soft ray always present ?); a scale-like spine and an internal rudimentary soft ray shorter than the scale. (12) The principal teeth of the premaxillary and dentary series are without obvious barbs and without special enamel thickenings at the tips. When barbs are present, usually on the premaxillary fangs, they are barely perceptible and confined to the hinder edges. The margins of the teeth are smooth in both jaws. (13) Palatine teeth present in a linear series, exposed (B. tenwis) or concealed under mucosa (B. simonyt). (14) Principal teeth on first gill-arch few, teeth becoming progressively reduced on subsequent arches. (15) Intermuscular bones (pleurals and epipleurals) reduced. (16) Pigmentation uniform silver sprinkled black. Melanophores thinly distributed, except for denser aggregations along lateral line and along median dorsal and ventral lines. Dark spots at bases of dorsal and anal rays, preceded by large individual stellate melanophores in juveniles. Fins shaded with pastel colours. ZOOL, 4, 3. ai 88 THE FAMILY TRICHIURIDAE Key to Species Ventral fins inserted before anterior end of pectoral base. Dorsal rays 120-133; anal elements 1 + I + 70-76 with external rays through- out ; vertebrae 123-131 ; lateral line strongly developed (less than 15 times in height at pectoral) Benthodesmus tenuis (Giinther) E. Equatorial Atlantic ; Gulf of Mexico ; Indo-Pacific. Ventral fins inserted behind posterior end of pectoral base. Dorsal rays 147-155; anal elements i + I + 91-99 with external rays substan- tially confined to posterior third; vertebrae 153-158; lateral line less strongly developed (more than 20 times in height at pectoral) Benthodesmus elongatus (Clarke) New Zealand ; Australia; S. E. Africa (?) Benthodesmus simonyi (Steindachner) N. Atlantic ; N.E. Pacific For full discussion and complete bibliographies see :— Tucker, 1953, Proc. zool. Soc. Lond. 123 : 171-197, pls. and text-figs. 1955, Bull. Mus. Hist. nat. Belg. 31, No 64: 1-26, 1 pl. and text figs. Benthodesmus elongatus (Clarke) Lepidopus caudatus (non Euphrasen, 1788) Hutton, 1872, Fishes of New Zealand : 13. Lepidopus elongatus Clarke, 1879, Tvans. N.Z. Inst. 11 : 294, pl. 14. Holotype should be in the Dominion Museum, Wellington, N.Z., but cannot be found (fide Mr. J. Moreland in litt.). Type locality Hokitika Beach, W. coast of South Island, New Zealand. Benthodesmus elongatus (part), Goode & Bean, 1882, Pyoc. U.S. Nat. Mus. 4: 380. Lepidopus (Benthodesmus) elongatus McCulloch, 1915, Biol Res. ‘“‘ Endeavour,” 3: 152. ? Benthodesmus atlanticus (non Goode & Bean, 1895) Gilchrist & Von Bonde, 1924, Rep. Fish. Mar. biol. Surv. S. Afr. 3, Spec. Rep. 7 : 16. ? Benthodesmus tenuis (non Giinther, 1877) J. L. B. Smith, 1949, Sea Fishes S. Africa : 312. Benthodesmus simonyi (Steindachner) (Text-fig. 8). ? Lepidopus elongatus Clarke, 1879, Tvans. N.Z. Inst. 11: 294, pl. 14. (See above.) Benthodesmus elongatus (part), Goode & Bean, 1882, Pyoc. U.S. Nat. Mus. 4 : 381. Aphanopus simonyi Steindachner, 1891, S.B. Akad. Wiss. Wien 100 : 356. Holotype should be in the Naturhistorisches Museum, Vienna, but cannot be found (fide Dr. D, Kahsbauer, in litt.). Type locality N.E. from S. Cruz de Teneriffe, Canary Is. Benthodesmus atlanticus (part) Goode & Bean, 1895, Oceanic Ichthyology : 206. Holotype U.S. Nat. Mus. Washington No. 29116. Type locality W. edge Grand Bank of Newfoundland. (The two smaller specimens mentioned are Diplospinus multistriatus Maul, fide Dr. Carl L. Hubbs in Jitt.) Lepidopus sp. Vieira, 1895, Ann. Sci. nat. Pério 1 : 165, upper figs. pl. 9 and ro. Lepidopus ailanticus, Boulenger, 1899, Ann. Mag. nat. Hist. (7) 3: 180. Lepidopus (Benthodesmus) atlanticus Saemudsson, 1921, Skyrsla um hide islenzka nattivufraedisf- jelag 1919-20 : 37. Benthodesmus tenuis (non Giinther, 1877) (part) J. L. B. Smith, 1949, Sea Fishes S. Africa : 312. (Figure copy of B. atlanticus from G. & B. 1895.) Benthodesmus simonyi Maul, 1953, Proc. zool. Soc. Lond. 123 : 167. THE FAMILY TRICHIURIDAE 89 Benthodesmus tenuis (Ginther) (Text-fig. 9) Lepidopus tenuis Giinther, 1877, Ann. Mag. nat. Hist. (4) 20 : 437. Lepidopus tenuis Giinther, 1887, ‘‘ Challenger ’’ Reps. Zool. 22 : 37, pl. 7, fig. B. Holotype B.M. (N.H.) No. 1879.5.14.297. Type locality “‘ Challenger’ St. 232, 35° 11’ o” N., 139° 28’ o” E., off Inosima, Sagami Bay, Japan. Benthodesmus tenuis, Goode & Bean, 1895, Oceanic Ichthyology : 206. Benthodesmus elongatus (non Clarke, 1879) idem. loc. cit. (figure only, a reversed tracing from Giinther, 1887). Lepidopus aomori Jordan & Snyder, tgot, J. Coll. Sci. Tokyo, 15 : 303. Holotype in the Aomori Museum, Japan. Type locality Aomori Bay. Benthodesmus benjamini (part) Fowler, 1938, Proc. U.S. Nat. Mus. 85: 45, fig. 16. Holotype U.S. Nat. Mus. No. 98821. Paratypes 98822-5. Type locality ‘“‘ Albatross ”’ St. D.5445, off Philippine Is. (The paratype material is contaminated with Diplospinus multistriatus Maul, fide Dr. Carl L. Hubbs, im litt.) Benthodesmus atlanticus (non Goode & Bean, 1895) Longley & Hildebrand, 1941, Cat. Fish. Tortugas : 73. ? Lepidopus argenteus (non Bonnaterre, 1788) Brauer, 1906, Wiss Evgebd. ‘“‘ Valdivia,” 15 : 292, taf. 12, fig. 3. (Fig. erroneously captioned L. gracilis.) Benthodesmus sp. incertae sedis Lepidopus tenuis (? non Giinther, 1877) Franz, 1910, Abh. Bayer. Akad. 4 Suppl. Bd. 1:56. (Locality Uraga Channel, Japan.) On the information available this specimen cannot be assigned with certainty to either B. simonyi or B. tenuis. I do not believe it to be a new species, nor do I accept Franz’s opinion that it justifies regarding this genus as containing one world-wide species. Subfamily LEPIDOPODINAE Gill Lepidopodinae Gill, 1863, Proc. Acad. nat. Sci. Philad. 1863 : 227. Type genus Lepidopus Gouan. GENERA NOW RECOGNISED.—Lepidopus Gouan; Evoxymetopon (Poey) Gill; Eupleurogrammus Gill; Assurger Whitley ; Tentoriceps Whitley. DIAGNOSIS : (Note.—Since there is considerable diversity among the genera of Lepidopodinae and since, through inadequate descriptions and lack of study-material, certain characters have not been verified in Evoxymetopon, Assurger and Tentoriceps, it is necessary to introduce qualifications into the following diagnosis. For this purpose the abbreviations Lep., Evox., Eupl. Ass., & Tent. have been used for the generic names). A. Slope of snout variable, gentle to steep; orbits barely entering upper profile of head (Lep.) or more or less remote from it (all other genera) ; posterior confluence of frontal ridges elevated to support a sagittal crest at the nape which (in all genera except Lep.) is continued forward along the snout as a ridge-like elevation of the entire ethmofrontal region. go THE FAMILY TRICHIURIDAE w Cartilaginous protuberance at mandibular symphysis weak or absent ; a small, soft projection at the tip of the snout. Lower hind margin of operculum markedly convex. | Teeth of main series without barbs. (Lep., Eupl., Evox., Tent. Ass.). (Fangs slightly barbed in Lep.). Teeth on palatines in a linear series. (Lep., Evox., Eupl.) Lateral line descending gently from the shoulder and running in a median or sub-median postiion along the body, i.e. distance between lateral line and ventral profile at anus much more than half distance between lateral line and dorsal. G. Spinous dorsal fin short, with 10 (Evox.), 9 (Lep.) or 3 (Eupl.) rays. Spinous and soft dorsals continuous, without any intervening notch. H. Soft dorsal rays precisely corresponding to adjacent caudal vertebrae, each basal and interneural element being related to a neural spine. (Lep., Ew.) I. Spinous anal i (Lep., Eupl.) + I (all genera) ; anterior soft anal rays not penetrating skin (Lep. Eupl. Evox. Ass.) and external and functional fin effectively confined to posterior ca. 20 rays, or (in Eupl.) absent. J. Terminations of dorsal and anal fins sub-opposite (Lep., Evox. Ass.) or anal extending alightly beyond dorsal (Ew). K. Caudal fin present, small, normal, forked (Lep., Evox., Ass.) or absent (Ewpl., Tent.) L. Ventral fins always present, composed each of a scale-like spine and some- times at least (Lep.) an internal rudimentary soft ray; insertion retarded, 1 to 5 eye-diameters behind posterior end of pectoral base. M. Pyloric caeca ca. 24 (Lep., Eupl.) mee SSS Osteological literature Giinther, 1860, Cat. Fish. B.M. 2 : 345-346 (short desc. Lepidopus). Starks, 1911, Stanford Univ. Publ. 5 : 17-26, pl. (skull of Lepidopus). Tucker, 1953, Proc. zool. Soc. Lond. 123 : 196, pls. (paired fins and anal of Lepidopus). Literature on young stages Delsman, 1927, Treubia 9, Livr. 4: 338 (Eupleurogrammus eggs and larvae). Regan, 1916, Sci. Rep. Brit. Antarct. Exped. Zool. 1: 144, pl. 8 (young Lepidopus). Strubberg, 1918, Rep. Dan. Oceanogr. Exped. 2 Biol. A.6.11 : 7-16 (life-history of Lepidopus). Genus LEPIDOPUS Gouan Lepidopus Gouan, 1770, Hist. Pisciwm : 107, 185, Tab. 1, fig. 4. No type species designated. Two species. } The earliest available binomen is L. arvgenteus Bonnaterre, 1788, Encycl. Méth. Zool. Icht. : 58, pl. 87, fig. 364. Bonnaterre’s figure is an accurate reversed tracing of Gouan’s caricature, but L. avgenteus is a synonym, and almost certainly a junior synonym, of Trichiurus caudatus Euphrasen, 1788, Handl. K. Vetensk. Akad. Stockholm 9: 52, tab. 9, fig. 2. Euphrasen’s paper appears in the section of the Handl. K. Vetensk. Akad. for Jan., Feb., Mar., 1788, the sections having been issued quarterly with separate title-pages though paginated in annual volumes. THE FAMILY TRICHIURIDAE y rE TTLEETTTMTETTTLLILICOUTILE FP 92 THE FAMILY TRICHIURIDAE In their third and final attempt to establish the dates of publication of the parts of the Encyclopédie Méthodique, Sherborn & Woodward, 1906, Ann. Mag. nat. Hist. (7) 17: 578 could establish nothing more precise concerning Bonnaterre’s Ichtyologie than that it appeared in livraison 28 of the Encyclopédie issued sometime in 1788, Since, however, the livraisons were issued in order and the date April, 1788, can be assigned to livraison 26 i the balance of probability favours Euphrasen’s publication as the earlier one. Following the nomenclatorial orgy at the earlier part of the nineteenth century Euphrasen’s name has been the more generally used. Synonyms (The full references to the following are given in the synonymy of Lepidopus caudatus, p. 93). Trichiurus (non Linnaeus, 1758) Vandelli, 1797 ; Holten, 1802. Vandellius Shaw, 1803. Type species Vandellius lusitanicus Shaw (ex Vandelli MS.). Ziphotheca Montagu, 1809. Type species Ziphotheca tetradens Montagu. Xiphotheca Xipotheca (Non Zyphotheca Swainson, 1839.) ? Scaycina Rafinesque, 1810. Type species Scarcina argyrea Rafinesque. DiaGnosis (based on L. caudatus) : (1) Body elongate, head 5-8-7-r in standard length, greatest depth 10-8-18-3 in standard length (57-1224 mm.) (2) Upper profile of head oblique-concave, rising at about 25° to the longitudinal axis from above the snout tip to behind the orbits and thereafter more steeply to the dorsal origin; straight before the orbits. Ethmo-frontal region not elevated, posterior confluence of frontal crests strongly elevated. Interorbital slightly concave with very low longitudinal ridges. (3) Orbit large, 4-9-5-6 in head, touching dorsal profile. (4) Dorsal IX, 90-96 ; aggregate 99-105. The first dorsal spine is not enlarged, save as a transient larval character. (5) Anal spinesi + I; Iisa small triangular scale 2 or more in the pupil. (6) Anal fin elements i+ 1I-+ 61-64; anterior rays reduced or absent, posterior 20-24 rays supporting fin. (7) Posterior end of operculum a broadly rounded point, barely reaching to anterior end of pectoral base. (8) Ventral fins present, scale-like, inserted an eye-diameter behind the posterior end of the pectoral base. (9) Caudal fin present. (10) Vertebrae 41 + 70-73 = III-113. |wariant spellings by later authors. Key to Species Dorsal rays 99-105 ; external anal fin reaching only half way to vent. D.IX, 90-96; analelementsi + I + 61-64 (the last 20-24 only being external fin-supporting rays) ; vertebrae 41 + 70-73. Head 5-8-7-1 instandard length 57-1224 mm. ; depth 14:4 (—18-3)—-10-8 insame ; eye 4°9-5°6 in head. Ventral fins I-1 (1 is an internal rudiment only 1 mm. long in the adult fish), inserted an eye-diameter behind the pectoral base ; anal spine I is a small triangular scale, less that the pupil. Pyloric caeca 20 +. Colour uniform silvery. Lepidopus caudatus (Euphrasen). Atlantic, Mediterranean, S. Indian Ocean, S, Pacific, THE FAMILY TRICHIURIDAE 93 Dorsal rays 82-87 ; external anal fin reaching to vent. Analysis of dorsal spines and rays not known; external anal i + I + 45-58; vertebrae unknown. For body proportions see discussion. Ventral fins I-1 inserted on or immediately behind perpendicular through posterior end of pectoral base ; anal spine I is long, keeled, about three-quarters the diameter of the eye. Pyloric caeca unknown. This compromise description, based on Jordan & Evermann (1898) and Brauer (1906), may include two species or one, of uncertain systematic position, and without valid name(s). The discussion on pp. 95-7 explains this unhappy situation. “ Lepidopus Xantusi’’ Goode & Bean California, Gulf of Guinea. Lepidopus caudatus (Euphrasen) (Text-fig. Io). Trichiurus caudatus Euphrasen, 1788, Handl. K. Vetensk. Akad. Stockholm, 9 : 52, tab. 9, fig. 2. Holotype in Alstromerika Museum ? Type locality Cape of Good Hope. Lepidopus caudatus, White, 1851, List Brit. Anim. B.M. 8 Fish: 32. Lepidopus argenteus Bonnaterre, 1788, Encycl. Méth. Zool. Ichth. : 58, pl. 87, fig. 364. (Ex Gouan, 1770.) (See note under Lepidopus p. 90.) There is a partial confusion with Lepturus argenteus Linnaeus, 1754 (= Trichiurus) in the text. Trichiurus ensifoymis Vandelli, 1797, Mém. Acad. Sci. Lisboa, 1: 70 (nomen nudum). (id. fide Nobre, 1935, Faun. Mar. Portugal, 1 Vert. : 260). Lepidopus ensifoymis, Swainson, 1839, Lard. Cab. Cycl. Fish. 2 : 254. Lepidopus gouanianus Lacépéde, 1800, Hist. nat. Poissons 2 : 519. (Ex Gouan, 1770.) Lepidopus gouani Bloch & Schneider, 1801, Syst. Ichth. 1 : 239, tab. 53, lower fig. (Ex Gouan, 1770.) Trichiurus gladius Holten, 1802, Sky. nat.-Selsk. Kbh. 5, Heft 2 : 23, Tab. 2, fig. 1. Holotype in Copenhagen Museum ? Type locality Portugal. (I am doubtful whether this name should not perhaps be attributed to Abildgaard.) Vandellius lusitanicus Shaw, 1803, Gen. Zool. Pisc. 4 (2) : 199. (Ex Vandelli MS.) Lepidopus lusitanicus, Leach, 1815, Zool. Misc. 2: 7, pl. 62. Ziphotheca tetradens Montagu, 1809, Mem. Werner. N. H. Soc. 1: 81. Holotype B.M. (N.H.) No. 1955.6.2.1. Type locality English Coast. Lepidopus tetvadens, Fleming, 1828, Hist. Brit. Anim. : 205. Lepidopus peronii Risso, 1810, Ichth. Nice : 148, Pl. 5, fig. 18. Type locality Nice. ? Scarcina argyrea Rafinesque, 1810, Car. n. gen. : 20, pl. 7, fig. I. Type locality Sicily. ? Lepidopus argyreus, Cuvier, 1829, Régne Animal 2 Ed. 2: 217. Lepidopus govanianus Risso, 1826, Hist. Nat. 3 : 290. (Ex Gouan, 1770.) Lepidopus lex Phillips, 1932, N.Z. Journ. Tech. 13 : 232. Syntypes in Dominion Museum, Wellington ? Type locality New Zealand. (Lepidopus caudatus of other New Zealand authors ; non L. caudatus Hutton, 1872, Fishes N.Z. : 13, who had Benthodesmus elongatus (Clarke).) Aphanopus carbo (non Lowe, 1839) (part) Norman, 1937, in Fraser and Norman, Giant Fishes, Whales and Dolphins : 140. non Lepidopus caudatus Sim, 1898, Ann. Scot. nat. Hist. 1898 : 53 (mis-identification of Aphanopus carbo Lowe). non Lepidopus elongatus Clarke (1879) ; McCulloch (1915) (see Benthodesmus elongatus). 94 THE FAMILY TRICHIURIDAE non Lepidopus sp. Vieira (1895). : : non Lepidopus atlanticus, Boulenger (1899) ; Saemundsson (1921) eS coe) non Lepidopus aomori Jordan & Snyder (1901) : non Lepidopus argenteus Brauer (1906) AB ee pede aaa) The nineteenth century synonyms listed above have been pretty generally accepted; Thave verified each of them, so far as the accompanying data allow, and do not propose to attempt any individual justifications in the present short summary. Only Scarcina argyrea Rafinesque (1810) calls for any urgent comment. This name has been copied as a synonym of Lepidopus caudatus by many authors, but the figure shows a head and body-form very reminiscent of a Benthodesmus and the stated dorsal count (125) falls within the range of B. tenwis (Giinther) and is well above the D.gg—105 found in L. caudatus. The anal count of 15 and the anal fin as figured are, however, quite like Lepidopus. Since Benthodesmus is not yet known from the Mediterranean it is better to regard Rafinesque’s as an inaccurate impression of L. caudatus for the present. Should B. tenwis be found in the Mediterranean Scarcina argyrea will have to be considered as a senior synonym and it may be thought desirable to invoke the Plenary Powers of the International Commission in order to suppress it. Scarcina would also precede Benthodesmus. Norman (1937) mentions Afhanopus carbo as being “‘ not uncommon in the fish markets of the Mediterranean ’’. In an intensive study of A. carbo I have so far found nothing to confirm this statement, which may have been made through some confusion of vernacular names. Thus the Portuguese and Madeiran fishermen call A. carbo “O Peixe Espada preta”’ and L. caudatus ‘“‘O Peixe Espada branca”’ (Black and White Scabbard-fishes, respectively), and in both cases “‘ Peixe-espada ”’ or“ Espada” for short. Phillipps (1932) : “examined several frost-fish and found consistent, though slight, differences between the New Zealand and Atlantic species . . . 3 to 4 less rays in the dorsal fin, 3 or 4 less anal rays, and a total length of head under 7 in total length. Goode and Bean’s figure shows a species with a longer head, and no dorsal spines of greater length than the diameter of the eye. In the New Zealand fish the height of the sixth dorsal ray is 5 in the head while in the European fish the height of this ray is about 8 or more in the head. The tail of the New Zealand frost-fish is not so deeply emarginate and agrees more nearly with that of Evoxymetopon taeniatus figured by Goode and Bean.” Phillipps is presumably referring to Goode & Bean (1895) Oceanic Ichthyology, Plate 58, figs. 213 and 214. I have dealt in some detail with the identification of fig. 213 under “ Lepidopus Xantusi” (p. 96 q.v.) and so for the present it is sufficient to state that this figure is a poorish figure of an apparent young Lepidopus caudatus and not a very satisfactory basis for any comparison. The head in Goode & Bean’s figure goes about 7-5 in the total length and is therefore shorter, not longer as stated by Phillipps, and typical of a juvenile as opposed both to post-larval and adult specimens. The dorsal spines and tail of Goode & Bean’s figure are useless as evidence, THE FAMILY TRICHIURIDAE 95 Comparing specimens of as nearly equivalent size as possible I obtain the following results : TABLE II. Lepidopus caudatus. Lisbon. New Zealand. No. 1860.4.22.69. No. 1903.4.30.29. Standard length . : 2 1142 mm. . 1224 mm. Head in S.L. 3 : : 7°13 ‘i 6°65 Depth in S.L. 2 : : 15°43 : 10°83 Eye in head length 5 : 4°92 5 5°41 6th dorsal spine in H.L. 0 5°33 c 5°25 Dorsal count 3 : D.IX, 96 D.IX, 90 Anal count . 0 c A.i+I+40+24 ° A.i+I+41+20 Apart from the greater depth of the body, in part attributable to age, the New Zealand specimen appears to show only the trivial differences to be expected in material of a widely ranging pelagic fish taken from the extreme limits of its distribu- tion. The variations are no greater than those found in Trichiurid species, of which I have been able to study substantial samples and accordingly I am not at present prepared to accept Lepidopus lex Phillipps as distinct from L. caudatus (Euphrasen). If, however, the separation of L. /ex should be considered justified an interesting situation arises. Since the type locality of L. caudatus is the Cape of Good Hope it is likely that the antipodal forms will be conspecific but distinct from those of the E. Atlantic and Mediterranean. L. lex would therefore still fall as a synonym of L. caudatus, but L. argenteus Bonnaterre (1788) would have to be revived. A further complication would arise in that the Lepidopus caudatus figured by Goode & Bean (r895) and uncertainly associated with the holotype of L. xantusi G. & B. appears to have the lower dorsal count of L. Jex also. (Further discussion under L. xantusz, p- 96). “Lepidopus xantusi” Goode & Bean Lepidopus caudatus (? non Euphrasen) Jordan & Gilbert, 1882, Proc. U.S. Nat. Mus. 5 : 358. Lepidopus caudatus (? non Euphrasen) (part) Goode & Bean, 1895, Oceanic Ichth. : 203, (?) fig. 213. Be eropas caudatus (? non Euphrasen) Jordan & Evermann, 1896, Bull. U.S. Nat. Mus. No. 47 : 886. Lepidopus caudatus (? non Euphrasen) Jordan & Evermann, 1900, Bull. U.S. Nat. Mus. No. 47, (°) pl. 136, fig. 373. Lepidopus xantusi Goode & Bean, 1895, Oceanic Ichth. : 519. Holotype U.S. Nat. Mus. No. to115. Type locality Cape San Lucas, California. Lepidopus xantusi Jordan & Evermann, 1898, Bull. U.S. Nat. Mus. No. 47 : 2843. Lepidopus xantusi Jordan & McGregor, 1899, Rep. U.S. Fish. Comm. 24 (1898) : 276. Lepidopus xantusi ? Brauer, 1906, Wiss. Evgebd. ‘‘ Valdivia,” 15 : 291, taf. 12, fig. 2. The circumstances surrounding the publication of this species are so wretchedly unsatisfactory that a new name will have to be found for it by the first worker able to re-describe it from material. Jordan & Gilbert (1882) list U.S. Nat. Mus. No. ro115, ‘“‘ One specimen, ro inches long, in poor condition ” as “‘ Lepidopus caudatus (Euphr.) White ” in a catalogue of 96 THE FAMILY TRICHIURIDAE the fishes collected by one John Xantus at Cape San Lucas, California. (The reader should beware confusion with the Joanne Xantus whose Asian collections were pub- lished by Karoli.) Goode & Bean (1895 : 203) give a description of L. caudatus evidently taken from Giinther whose name is, in fact, cited. They then refer toa Xantus specimen and on p. 13 of the accompanying Aflas of plates they state that their figure of L. caudatus is drawn from U.S. Nat. Mus. No. ro115, collected by John Xantus, off Cape St. Lucas. On p. 519 of an appendix to the main text, however, this specimen becomes the type of a new species with the brief remark : “ The specific identity of the fish found at St. Lucas by Xantus is so doubtful that we prefer to refer to it as L. Xantusi, new specific name.” We are left to consider whether Article 21 of the International Rules has been complied with ; on the text alone L. Xantusi is a nomen nudum and may be saved only by the figure, to be discussed presently. Jordan & Evermann (1896) give the Giinther-Goode & Bean version of L. caudatus (with an addition of pure Giinther) and conclude by assigning the Xantus specimen once again to L. caudatus. Jordan & Evermann (1898) have realized that L. Xantust exists and that somebody should give a description of it, but instead of describing it from the holotype (10 inches S.L.) they elect to do so from a second Cape San Lucas specimen which is more portable (54 inches S.L.). Jordan & Evermann (1900), however, continue to publish Goode & Bean’s original figure of the supposed holotype of L. Xantusi still with the legend “ L. caudatus” The figure published by Goode & Bean has no scale of magnification nor do these authors anywhere state the size of their specimen; for that we have to return to Jordan & Gilbert (1882). Moreover the drawing has the tail nicely curved, an effective obstruction to accurate measurement of standard length. I derive the following data : Radial Formula D.g9; A. (external) 18. (mm.) Measured distance from snout tip to D.30 . : 125 Estimated distance from D.30 to D.70 (taken as 4 x mean acters) D.20-30 and D.70-80) . 5 : “i = £20 Measured distance from D.70 to tip caudal ‘peduncle : c a ah) Whence’ Estimated standard length of figure . ° 3 > 5 6 See Head in S.L. 7:3; depth in S.L. 18-6; eye in head 5-7; snout in head 3:1 Insertion of ventral fins an eye-diameter behind pectoral base. But these are the counts found in Lepidopus caudatus (Euphrasen) and these the body-proportions of a young fish of that species! We are therefore driven to one of two conclusions : Either (1) The figure is drawn, by some accident, from a specimen other than the holotype of L. Xantusi Goode & Bean. In this case the name L. xantusi Goode & Bean falls as a synonym of L. caudatus (Euphrasen); whatever the tent THE FAMILY TRICHIURIDAE 97 identity of the Xantus specimen, no ‘‘definition or description ’’ have been pub- lished, nor any figure of that specimen. Further, although Jordan & Evermann (1898) and Brauer (1906) give adequate characterisations of a species distinct from L. caudatus (Euphrasen) under the name L. Xantusi, their name must fall as a homonym of L. Xantusi Goode & Bean under Article 35 of the Rules. or (2) The figure is drawn from U.S. Nat. Mus. No. rorrs as stated and represents the holotype of L. Xantusi G. & B. In this case L. Xantusi again falls as a synonym of L. caudatus (Euphrasen) and L. Xantusi Jordan & Evermann and L. Xantusi Brauer again fall, as homonyms, under Article 35 of the Rules. The description by Jordan & Evermann (1898) is repeated verbatim by Jordan & McGregor (1899). I give the complete text : “Head 4 2/3 in body; depth 3 in head; eye 5 1/3; interorbital space 8 1/3; snout 3; maxillary 3 1/3. D.82; A.II, 45. Jaws with long, sharp teeth in front, followed by single rows of weaker ones, arranged in groups of twos and threes. Height of dorsal, near middle of body, 3 in head. Anal preceded by 2 scutes, the first minute, the second wide, strongly keeled, its length 3/4 the diameter of eye. Pectorals of 12 rays, length 2 in head. Each ventral consists of a flat keeled spine followed by a minute ray. This species is known from 2 small mutilated specimens, both found on the beach near San Jose del Cabo, Cape San Lucas. The type was taken by John Xantus, about 1860, and recorded by Jordan & Gilbert as Lepidopus caudatus. The second, of about the same size (53 inches), was taken by Richard C. McGregor, in 1897. From the latter the above account was taken. The species differs from Lepidopus caudatus in the much shorter dorsal and longer anal. D.103 ; A. 24. (Named for John Xantus de Vesey).” Additional data, not provided above, are now needed to decide whether this fish may remain in Lefidopus when a new name shall be assigned to it ; at present it could as well belong to an Aphanopodine genus as to Lepidopus and may even represent a new genus connecting Diplospinus and Lepidopus. Brauer (1906) gives a description and figure of “ZL. Xantusi”’ from the Gulf of Guinea and discusses the difficulties of his identification in face of the above descrip- tion. The size is not given, but since a scale of magnification is given for some of the other figures on the same plate (though not for this) we may assume Xz, hence 1 5Imm. S.L. Brauer gives D.87; A.58; head 5-5 in S.L.; depth 15 in S.L. ; eye 54 in head. Tt would help if Jordan & Evermann meant “ Head 4 2/3 in body (less head) ”’, i.e. 5 2/3 inS.L., which would also give depth 17 in S.L. instead of 14. The discrepancies between the fin-ray counts are obvious. The figure shows a head about intermediate in form between Aphanopus and Lepidopus and ventrals inserted barely behind the pectorals, not quite so far retarded as in L. caudatus. Clearly we should know more about these specimens. Genus EVOX YMETOPON (Poey) Gill. Evoxymetopon Poey, in Gill, 1863, Proc. Acad. nat. Sci. Philad. 1863 : 227. Type species Evoxymetopon taeniatus (Poey) Gill. Monotypic, or two species, TRICHIURIDAE THE FAMILY 08 / aie (‘papper ayeos “Ty cur o1t'r ‘adAjyojoH CRESS :uevag 3 apoosy wo; ‘UMPIP-ay{) TED (Avog) snzoruan} uodojamAxon q— Il “SI A THE FAMILY TRICHIURIDAE 99 DIAGNOSIS : (1) Body elongate, head 8 in total length, greatest depth 12-13 in total length. (1410-1980 mm.) (2) Upper profile of head convex, a steep continuous curve from the tip of the snout to the origin of the dorsal set at about 45° to the longitudinal axis ; Slightly convex before the orbits. Structure of cranial crest un- known, but evidently the ethmo-frontal region and the posterior con- fluence of the frontal crests are both elevated. Interorbital strongly convex. (3) Orbit large, 5-6 in head length, an eye-diameter + below the dorsal profile. (4) Dorsal X, 77; aggregate 87. The first dorsal spine may be enlarged, nearly as long as the head. (5) Anal spines i(?) + I; Lisa keeled scale. (6) Analysis of anal fin elements unknown; the anterior rays, if present, appear barely to penetrate the skin while the posterior ca. 20 only are fin-supporting rays. (7) Posterior end of operculum a broadly rounded point falling less than a pectoral base short of the pectoral base. (8) Ventral fins present, scale-like, inserted an eye-diameter behind the posterior end of the pectoral base. (9) Caudal fin present. (10) Analysis of vertebrae unknown. Evoxymetopon taeniatus (Poey) Gill (Text-figs. 1 12 13) Evoxymetopon taeniatus Poey, in Gill, 1863, Pyoc. Acad. nat. Sci. Philad. 1863 : 228. Holotype U.S. Nat. Mus. No. 5735. Type locality Havana, Cuba. Evoxymetopon taeniatus Poey, 1873, Ann. Soc. Esp. Hist. nat. Madrid, 2:77, pl. 5. Evoxymetopon taeniatus Goode & Bean, 1895, Oceanic Ichthyology : 204, fig. 214. ? Evoxymetopon poeyi Giinther, 1887, ‘‘ Challenger’ Reps. Zool. 22 : 39, pl. 43. Disposal of holotype unknown. Type locality Mauritius. (For Evoxymetopon anzac Alexander see under Assurger, p. 106.) It is curious that Poey should have waited ten years before publishing his own description and first figure of this species. Goode & Bean give a new figure of the holotype but their description appears to be derived entirely from Gill and their only original contribution is to confuse Gill’s percentages with millimetres and so to mislead others into believing that the specimen is only one-fourteenth of its true length. Evoxymetopon poeyi, described “ with great hesitation . . . as a second species” was based on a dry skin which Giinther received from Mauritius while his “‘ Challenger’ Report was passing through the press. The ownership and ultimate destination of this specimen are unstated and unknown ; there is certainly no evidence that it ever became part of the permanent collections of the British Museum (Natural History). The salient characters of these two fishes, as compiled from the literature, are given in Table III, from which it is apparent that there is a large measure of agreement between them and that the differences are readily attributable to age or sex, damage or misinterpretation. 100 THE FAMILY TRICHIURIDAE The elongated first dorsal spine noted in E. foey? is a striking feature and apparently unique among adult Trichiurids ; though it occurs in the young stages of Lepidopus it does so only as a transitory condition and one to be regarded, like the disproportion- ate ventral fins, as a flotation device, parallelled among many other young Teleosts and without phyletic significance. E. taeniatus and E. poeyi may be female and male of Fic. 12.—Evoxymetopon taeniatus (Poey) Gill. Head of holotype, 1,410 mm. T.L. (from Poey). — 10 CM. Fic. 13.—Evoxymetopon poeyi Giinther. Head of holotype, 1,980 mm. S.L. (re-drawn, after Giinther ; scale added). one species (c.f. Anthias) or there may be growth changes between 1410 and 1981 mm. length, but a quite likely explanation is that the Cuban specimen may be damaged. The homologies of the parts in other Trichiuridae studied indicate that the post-anal structures probably comprise the usual minute spinule (so far overlooked) and a THE FAMILY TRICHIURIDAE IOI broad scale having a median depression or keel, the pair articulating with a simple or compound basal structure. Experience with damaged Benthodesmus material provides a ready explanation of how the discrepancies between the accounts of Gill, Poey and Giinther may have arisen. Total length Greatest height/T.L. Head length/T.L. Orbit/head Dorsal rays First dorsal spine Anal rays . Post-anal scute . Ventral insertions Vent Coloration TABLE III. Evoxymetopon taeniatus (Poey) Gill. 1410 mm. (Poey) 1/12 (Gill) 1/8 (Gill) 1/6 (Gill) X, 77 (D.87. “The first ten dorsal spines are undivided ; the rest split.’”’—Gill.) No special mention in either Gill or Poey. A.1g. “ Anal spines numerous - mostly minute, free, pos- teriorly enlarged, connected by the membrane and forming a fin ’’ (Gill). Upwards of 30 small spines figured anterior to the fin proper (Goode & Bean). “ Dagger-shaped spine behind the anus ”’ (Gill). “A corta distancia posterior del ano la pequefia escama trian- gular y movediza senelada por Cuvier en el Lepidopo”’ (Poey). About 14 times the head length from the tip of the snout (17} : 12—Gill). “ Submedian ”’ (Gill). “ Silvery, with about six narrow reddish bands most distinct behind, the first on the ridge of the back and the fifth along the lateral line ’’ (Gill). Evoxy metopon poeyi Giinther. 78 inches (Giinther) (ca. 1981 mm.) 1/13 or less (Giinther) 1/8 (Giinther) 1/5 (Giinther) D.93 (Giinther) c large, compressed, sword-shaped . . . not much shorter than the head loosely articulated with the interneural ’’ (Giinther). x + 20 (Giinther) - anal fin, the rays of which begin to be free in the posterior third of its extent ’’ (Giinther). (Gill’s spine) “‘ is entirely cover- ed by skin, and consists of coalesced and flattened inter- haemalelements . . . a single oval scale slightly bent along the middle occupies the space at a short distance behind the vent ’’ (Giinther). About 14 times the head length from the tip of the snout (313: 240—Giinther’s fig.). “Somewhat in advance of the middle of the total length” (Giinther). “Uniform _ silvery ” (Giinther). 102 THE FAMILY TRICHIURIDAE The upper profile of the head in E. poeyi does not rise quite as steeply as in E. taentatus (a condition apparently related to allometric growth of the jaws) and the whole head is less plump in appearance. Here, again, one recalls post-mortem changes witnessed in freshly caught Aphanopus carbo off Madeira, as well as the fact that E. poeyt is figured from a dry skin, and accordingly one discounts the differences. Gill (1863) alludes to a Scottish specimen referred by Hoy to Trichiurus lepturus (there were actually two) and suggests that it may have been an Evoxymetopon. Evoxymetopon has never been taken in British waters and Hoy’s specimens must be referred probably to Tvachypterus or Regalecus. Although the osteology of Evoxymetopon is unknown it is certain that the ethmo- frontal region of the skull, together with the posterior confluence of the frontal ridges must be elevated in much the same way as in Eupleurogrammus, but to a greater extent. This character apart Evoxymetopon stands fairly close to Lepidopus and is very near to the ancestor of Euplewrogrammus. Genus EUPLEUROGRAMMUS Gill Eupleurogrammus Gill, 1863, Proc. Acad. nat. Sci. Philad. 1863 : 226. Type-species Tvichiurus muticus Gray. Two species. Trichiurus (part). Many authors, from Linnaeus (1758), whose type material of Trichiurus leptuyus was contaminated with this genus. (See note under Tvichiurus, p. 114). Enchelyopus (part) Bleeker, 1872, Ned. Tijdschr. Dierk. 4 (1872) : 131. DIAGNOSIS : (r) Body very elongate, head 9-4-11-2 in total length, greatest depth (in region of vent) 14:7—-17-6 in total length (273-617 mm.) (2) Upper profile of head oblique to very slightly concave, rising from the tip of the snout in a line set at about 30° to the longitudinal axis and quite straight before the orbits. Cranial crest formed by elevation of ethmo- frontal region and of the posterior confluence of the frontal crests. Inter- orbital strongly convex. (3) Orbit small, 6-0-7-8 in head, $ to + an eye-diameter below the dorsal profile. (4) Dorsal III, 123-131 or III, 143-147; aggregate 126-150. First dorsal spine not enlarged. (5) Anal spinesi +1; [isa small triangular scale. (6) Anal fin elements 1 + I + 114-121; the external fin is entirely suppressed and the ventral profile smooth. (7) Posterior end of operculum a rounded point, overlying middle of pectoral fin and base. (8) Ventral fins present, scale-like, inserted about 5 eye-diameters behind the posterior end of the pectoral base. (9) Caudal fin absent. (10) Vertebrae 32-35 ++ 125-128 = 157-162 or 4I + 150-15I = I9I-192. THE FAMILY TRICHIURIDAE 103 Key to Stecies. Anal origin below D.33-37 D.III, 123-131; Vertebrae 32-35 + 125-128 = 157-162 Eupleurogrammus intermedius (Gray) Indo-Pacific. Anal origin below D.41-42 D.III, 143-147; Vertebrae 41 + 150-151 = 191-192 Eupleurogrammus muticus (Gray) Indo-Pacific. Eupleurogrammus intermedius (Gray) (Text-fig. 14) Trichiurus intermedius Gray, 1831, Zoo. Misc. 1: to. Syn-types (3) B.M. (N.H.) No. 1869.3.19.76. Type locality Chusan. Trichiurus muticus (non Gray) numerous authors. (Incorrect deductions from Gray's original description or from following Giinther, 1860, Cat. Fish. B.M. 2:348; no new material involved.) Trichiurus medius Griffith, 1834, Cuvier’s Anim. Kingd. Pisces: 349 (nom. emend. from Gray). Trichiurus savala (non Cuvier, 1829) (part) Bleeker, 1852, Verh. Bat. Gen. 24 Makr.: 41. (Deter- mination altered to T. glossodon by Bleeker, see below.) Trichiurus glossodon Bleeker, 1860, Acta. Soc. Indo-Neerl. 8. Dertiende Bijdr. Visch. Borneo : 38. ? Syn-types, in Leiden Museum and in British Museum (Natural History), B.M. (N.H.) No. 1880.4.21.119. Type localities Java, Sumatra, Singapore, Bintang, Borneo. Trichiurus glossodon De Beaufort, 1951, Fish. Indo-Austr. Aychip. 9: 190. (Bleeker’s material re-examined.) Trichiurus glossodon Delsman, 1927, Tveubia 9, livr. 4 : 338. Giinther (1860) regarded Tvichiurus intermedius Gray as a synonym of T. muticus Gray. This error of judgment not only led almost every subsequent worker astray ; it also had the practical result that Gray’s types in the British Museum (Natural History) were not properly recognized and segregated. There is, however, one jar, Reg. No. 1860.19.76, containing three specimens and bearing (among others) a label in an old hand stating : “ Trichiurus intermedius Chusan. E. I. Company.” A second label, written in ink on paint, changes the identification to Trichiurus muticus and a third, overlying both, adds the Register number and changes the source to “ Dr. Cantor’s Colln.” It is not possible to reconcile this material with any entry in Giinther (1860), but there seems no doubt that these are the syntypes of T. intermedius Gray, both from their apparent history and their study. Accordingly T. intermedius provides one of the major nomenclatorial surprises of the present paper. Even as a synonym of T. muticus it would, of course, have passed over into Eupleurogrammus, but, as shown in the key and in Table IV. T. intermedius proves to be a perfectly valid species. Further, on the evidence of a probable syntype of T. g/ossodon Bleeker and on De Beaufort’s (1951) re-description of other presumed type-material of T. glossodon at Leiden, it becomes apparent that the more widely-recognized T. glossodon is only a synonym of T. intermedius, as ZOOL. 4, 3. 8 THE FAMILY TRICHIURIDAE “T'S ‘wu Off Jo uemtoads ‘9 jo suowtoads yosproy “gq “yw ‘sedAquAs “(AvI5) ot: snip Ss wu goe ‘ snu €lz AUDASOANAIGnA— FI “OL THE FAMILY TRICHIURIDAE 105 TABLE IV. A. Standard Head Depth Eye origin Length in in in Fin-ray counts : below Vertebral (mm:)) S:E; S.L. head. D. A. DS} counts. Eupleurogvammus intermedius : Trichiurus intermedius Gray Syn-types: Chusan: East ( 273 . 10:11 . 17°61 . 6:00. III, 130 i+I+120 . 35-6 . 34+128=162 India Co. Reg. No. B.M. 308 = O-77. 17-tr . 6-56.) TIT, m28) G--E-Pr16),. 37) - 35-4-127—162 (N.H.) 1860.3.19.76 il 336 . To-29 «. 16:39 . 6:56. TIT, 13m i--T--114 - 35 - 32-+-127=159 Trichiurus glossodon Bleeker ? Syn-type: No. loc.: Bleeker Coll. Reg. No. B.M. (NE) 1880.4.21.119 E 427 . 9-47 . 15-00 . 6-42). III; 123 1--I--104 . 433) = 32--125—157 Eupleurogrammus muticus : Trichiurus muticus Gray Holotype: India: Hard- wicke, Reg. No. B.M. (N. eal ca. ca. 1955-5-13-2 - a 426 . 10°26 . 14:69 . 7:0 . III, 143 i+1+120 . 41 . 41-+151=—192 Eupleuvogrammus muticus (Gray) No data. Reg. No. B.M. (N.H.) 1955.6.4.1 . 2 (657 3 \EES2E T5642. 7855) (LUE t47) 1 L-ret,s 42 = 4r--r50—To0 Bleeker (1860) himself suspected, and Delsman (1927). Delsman goes far towards recognizing the different vertebral counts in 7. muticus and T. glossodon, but in the former case he appears to have had the misfortune to select a specimen with ; a broken tail and gives 40-115 = 155. De Beaufort comprehends the affinities of T. muticus and T. glossodon, as he shows in his key, but. apparently not having heard of Eupleurogrammus, he retains both species among Tvichiurus and gives no Eupleuro- gvammus combinations in his synonymies. De Beaufort’s counts on the type material of T. glossodon (D.115-120. A. about go.) seem a little on the low side though, in the nature of the material, not disturbingly so. Discussion of the relationships of the two species of Euplewrogrammus and of their systematic position is continued under the following species. Eupleurogrammus muticus (Gray) (Text-fig. 15) Trichiuvus leptuyus (part) Linnaeus, 1758, Syst. Nat. Ed. 10 : 246 (fide Lonnberg et al., 1896, K. Svensk. Vet.-Akad. Handl. 22:40. See note under Tvichiurus, p. 114.) Trichiurus muticus Gray, 1831, Zool. Misc. 1: to Holotype B.M. (N.H.) No. 1955.5.13-2. Type locality India. Eupleurogrammus muticus, Gill, 1863, Proc. Acad. nat. Sci. Philad. 1863 : 226. Enchelyopus muticus, Bleeker, 1872, Ned. Tijdschr. 4 : 131. non Trichiurus intermedius Gray, 1831, Zool. Misc. 1: 10 The taxonomy of Eufleurogrammus muticus requires very little comment, except a further emphasis on the fact that T. intermedius Gray is not a synonym of it as Giinther has led too many to suppose. I have examined the types of both nominal species and give the results in Table IV. The holotype of E. muticus has the gill-covers partly damaged, but another and better specimen has been available also. 106 THE FAMILY TRICHIURIDAE It has been discussed whether the distinction holds that E. muticus is “* burnished silver” and E. intermedius “ purely silvery’; Delsman and Day pro, De Beaufort con. De Beaufort appears to clinch the matter when he says : “My specimens” (of muticus) “do not differ in colour from specimens of glossodon ‘(= intermedius)’ collected in the same locality and preserved in the same jar.” The type of silver coloration is, in fact, quite variable in any one species of Trichiurid, depending on age, on fixative and preservative, on the amount of oil in the tissues (which can import a golden tinge to the silver) and on the fine or coarse grain of the guanine itself. Despite the superficial resemblance and absence of a caudal fin it seems surprising that these two species should ever have been placed in Tyvichiurus and still more so that Gill (1863) should have been content to recognize Eupleurogrammus without removing it to the Lepidopodinae. The typically Lepidopodine palatine teeth and median lateral line are fundamentally different from those of the Trichiurinae ; to these characters are allied a rounded operculum and the presence of ventral fins. Further, though the development of the cranial crests is unlike that of Lepidopus as of the Trichiurinae, it is very like that of Evoxymetopon, a Lepidopodine which Gill had in his hands and classified as such. The dentition of the main series is finer than that of any other genus of the Trichiuridae. Authors have regarded the ecaudate genera as “‘ degenerate ’’ simply because of their lack of a caudal fin. This is a very hasty and unwise opinion : in fact Ewpleuro- gvammus is one of the most advanced. Not only does it display the culminations of a number of progressive trends (see pp. 125-8); in appearance and structure it has the most elegantly streamlined form. The sides of the head and operculum are smoothly curved, with none of that chunkyness which occurs in the more primitive genera ; the upper and lower profiles of the body are both gently convex ; the dorsal is arched and the always untidy anal entirely suppressed ; the point of greatest depth has moved back toward the vent ; and a comparison of a radiograph of the skeleton with that of, say Nesiarchus or Diplospinus, is like a comparison between the mecha- nism of a high-grade chronometer and of a cheap alarm-clock. Genus ASSURGER Whitley Assurgey Whitley, 1933, Rec. Aust. Mus. 19: 84. Type species Evoxymetofon anzac Alexander. Monotypic, Indo-Pacific. DIAGNOSIS : (1) Body extremely elongate, head 12 in total length, greatest depth 28 in total length (ca. 1415 mm.) (2) Upper profile of head oblique, rising continuously from the tip of the snout in a straight line set at about 25° to the longitudinal axis and quite straight before the orbits. Structure of cranial crest unknown, but evidently the ethmo-frontal region and the posterior confluence of the frontal crests are both elevated. Interorbital strongly convex. (3) Orbit small, 8 in head length, 4 an eye-diameter below the dorsal profile THE FAMILY TRICHIURIDAE 107 (4) Analysis of dorsal spines and soft rays unknown; aggregate ca. 120. First dorsal spine not enlarged. (5) Anal spines i(?) + 1; Lisa large oval scale. (6) Analysis of anal fin elements unknown ; only the posterior 14-15 appear to be external fin-supporting rays. (7) Posterior end of operculum a rounded rectangle, falling about a pectoral base-length short of the pectoral base. (8) Ventral fins present, scale-like, inserted about 14 eye-diameters behind the posterior end of pectoral base. (9) Caudal fin present. (10) Analysis of vertebrae unknown. Assurger anzac (Alexander) (Text-fig. 16 and PI. ro). Evoxymetopon anzac Alexander, 1916, J. Roy. Soc. W. Aust. 2: 104, pl. 7. Holotype in the Western Australian Museum, Perth. Type locality North Fremantle, Western Australia. Evoxymetopon anzac Kamohara, 1952, Sci. Rep. Kéchi Univ. No. 3: 31, fig. 26. Assurger alexanderi (‘‘nom. emend., as Anzac is not permissible ’’) Whitley, 1933 ,Rec. Aust. Mus. 19: 84. (Whitley’s emendation is quite unnecessary since a ‘‘ Recommendation ”’ at the end of Article 14 of the “ International Rules of Zoological Nomenclature’ expressly states : “Latinized Greek words or barbarous words may, however, be used. Examples. . . giczac ...') This species is known from Alexander’s original and incomplete description and figure, the latter a photograph showing the head and the trunk back to the level of about the tenth dorsal ray. Whitley, in a general paper of miscellaneous studies, erected Assurger apparently on Alexander’s account alone and without examination of the specimen which, though remote from Sydney, must surely have been more easily accessible to Mr. Whitley than to any worker outside Australia. The following are all the data that can be extracted from Alexander : “IBS NOE edhe) TAO) PB HsNaval to (Cag) o 1B 57). Total length 1415 mm., length of head 120 mm., greatest height 50 mm., diameter of orbit 15 mm. “ Unfortunately the fins are a good deal broken, and it is impossible to count the rays of either the dorsal or anal with accuracy, no doubt these breakages occurred when it was washed ashore, and if the large spine at the commencement of the dorsal found in E. foeyi was ever present it has disappeared. In other respects the example agrees in its structural features with Giinther’s description, the postanal spine is exposed evidently owing to the abrasion of the skin in that region and just behind it there is a large oval scale similar to that described and figured by Giinther. There is no trace of the six narrow reddish bands which Poey describes in E. taeniatus and if one may judge from Goode and Bean’s figure, the ridge on the forehead is not nearly so high as in that species, but DD 4G agrees with that of E. poey?. ,., a bright silvery colour,” THE FAMILY TRICHIURIDAE 108 (peyono}-ar Aptavay ‘(zS6r) ereyourey JozFy) ‘uedef “JS ‘wut oSz‘z jo ueuedsS +9 (Iapuexe[y) IzuH Aadinssp—'9OI “OT , ae 2 = qj = CRMANSAAATANESESAASS. RS SSE THE FAMILY TRICHIURIDAE 109 Alexander’s references are to Poey (1873) and Goode & Bean (1895) (given with other Evoxymetopon references on p. 99). Although his discussion mentions Gill (1863) and Giinther (1887) also, there is internal evidence in his paper that he cannot have examined them all. Gill clearly states that the type of E. taeniatus is nearly five feet long (in which he is followed by Giinther) and Poey gives 1410 mm. total length, yet Alexander follows the mistake of Goode & Bean, who copy Gill’s percentage proportions of total length as millimetres, and gives Ioo mm. as the total length. When, therefore, Alexander denounces Goode & Bean’s “‘ very poor figure’’ of E, taeniatus he is complaining of a figure of a fish which he has not seen and which he has not compared with Poey’s independent drawing of the same specimen. Alexander’s figure shows the profile of the head rising in practically a straight line from the tip of the snout to behind the eye, the slope (with the mouth open) being about 25° to the longitudinal axis of the body. The eye lies half its diameter from the dorsal profile. The hinder end of the operculum falls about a pectoral base-length short of the pectoral fin. The ventral fins are not mentioned ; the photograph shows a nondescript median projection before the level of the end of the operculum, certainly irrelevant, and a slight indentation, about an eye-diameter behind the pectoral base, which is a likely position for the ventrals but quite inconclusive. Dr. L, Glauert, Director of the West Australian Museum, has kindly done what he could to amplify Alexander’s account. He has provided the original photographic print from which Alexander’s plate was made and this is reproduced, I hope with greater clarity than before, as Plate 10 of the present paper. One point which this print does clarify is the fact that there are no barbs on the teeth; Alexander’s plate may seem to indicate a barb on one of the premaxillary fangs, but this is an artifact due to indistinct reproduction of a piece of rubbish on the tooth in question. Dr. Glauert gives the eye-diameter as 16 mm. and the length of the head (measured from the snout-tip) as 113 mm., whence the ratio eye/head-length must be 1/7 instead of 1/8 as given by Alexander. Dr. Glauert is unable to add precision to Alexander’s account of the dorsal fin: “‘ . . . a University Undergraduate, interested in fishes, made an attempt and counted only 127, whereas all those others who made the attempt gave from 135 to 142. The explanation is that the dorsal fin was very much damaged when the fish reached the Museum ”’, Kamohara (1952) reports one specimen of 2250 mm. from Kéchi Market, Japan, and gives a small figure but no description. The illustration shows the fish bent into an S which prevents measurements of the body proportions and a count of the dorsal fin rays, but the general picture agrees with Alexander’s description and figure. One new fact emerges: the ventral fins are inserted about 1} eye-diameters behind the pectoral base. Since it is evident that the structure of the ethmoid and frontal region of the head must be similar to that in Eupleurogrammus and Evoxymetopon 1 place this fish among the Lepidopodinae, among which it may be considered to parallel Benthodesmus among the Aphanopodinae. It is at once separable from Eufleurogrammus and Tentoriceps by its possession of a caudal fin and from Lepidog¢us by the form of the head and of the elongate body. It differs from Evoxymetopon in the gentler slope of the snout, smaller eye (1/7: 1/6 of the head), elongate body (height 1/28: 1/12, head 110 THE FAMILY TRICHIURIDAE 1/12: 1/8 of total length) and higher dorsal count (ca. 120+ : 87). These compari- sons are between holotypes of practically identical size (Assuwrger anzac 1415 mm., Evoxymetopon taeniatus 1410 mm, total length). Genus TENTORICEPS Whitley Tentoriceps Whitley, 1948, Rec. Aust. Mus. 22: 94. Type species Tvichiurus cristatus Klunzinger. Monotypic. DIAGNOSIS : (1) Body extremely elongate, head 9 in total length, greatest depth 20-24 in total length (ca. 418 mm.) (2) Upper profile of head convex, a continuous curve rising from the tip of the snout at about 30° to the longitudinal axis and markedly convex before the orbits. Structure of cranial crest unknown, but evidently the ethmo- frontal region and the posterior confluence of the frontal crests are both elevated, the former perhaps disproportionately so. Interorbital convex. (3) Orbit large, 5 in head length (Kluzinger description) or 6 (Klunzinger figure), 2/3 of an eye-diameter below the dorsal profile. (4) Analysis of dorsal spines and soft rays unknown; aggregate ca. 120. First dorsal spine not enlarged. 5) Analysis of anal fin elements unknown : “ mit rudimentaren, kaum sicht- (6) baren Stachelchen ”’. (7) Posterior end of operculum acutely elliptical, reaching to middle of, but not concealing, pectoral base. 8) Ventral fins present, scale-like, but insertion unknown. 9) Caudal fin absent. 0) Analysis of vertebrae unknown. — Tentoriceps cristatus (Klunzinger) (Text-fig. 17). Trichiurus cristatus Klunzinger, 1884, Fische Rothen Meeres 1 : 120, Taf. 13, fig. 5a. Syntypes retained in Klunzinger’s private collection, Stuttgart; eventual disposal unknown. Type locality Kosseir, Red Sea coast of Egypt. Tentoriceps cristatus, Whitley, 1948, Rec. Aust. Mus. 22: 94. All that is known of this species is contained in Klunzinger’s original description and figure. I give the complete text : “ Kopfprofil convex, gratartig, scharf: eine hohe blattartige, bogige Crista zieht vom Beginn der Riickenflosse an tiber Stirn und Schnauze ; den vorderen Theil der letztern indess nicht mehr scharfend. Das Auge liegt daher weit unter der Profillinie. Bauchflossen wie beim vorigen in Form eines Schuppenpaares wie bei b.” (b. is Trichiurus muticus Gray, type-species of Eufleurogrammus.) “ Die Seitenlinie senkt sich sehr allméhlig abwarts und lauft etwas iiber dem unteren KG6rperdrittel. Afterflosse nur mit rudimentaren, kaum sichtbaren Stachelchen. Auge gross, 5 in der Kopflange, Schnauze yon doppelter Linge THE FAMILY TRICHIURIDAE Ill des Auges, Kopf massig lang, 2? mal so lang als der K6rper hoch, 9 in der Gesammtlange. Kérperhohe 20—24 (letzteres bei Aelteren) in der Gesammtlange, Korper also sehr gestreckt. Riickenstrahlen 1 1/4 in der Kérperhohe, 3 1/2 in der Kopflange, also ziemlich nieder. Brustflossen kurz, 7 in der Kopflange, (wenn nicht abgebrochen?). Peitsche kurz, nur von 1/2 Kopflaénge. Vordere Zahne einfach ohne Ansatz. D.c. 120 (?). Neue Art vom Rothen Meer.” “ Von dieser neuen und durch die scharfe, blattartige Kopfgrate gut charak- terisirten Art (siehe obige Uebersicht) bekam ich 3 Exemplare bei Koseir, ebenfalls aus dem inneren Meer. Farbe silbrig, Ruckenflosse hyalin ’’. The lengths of the three specimens are not given. The head, however, figured “‘ in nattirlicher Grésse”’ is 46-5 mm. in length, (measured from the snout tip) and this multiplied by 9 gives ca. 418-5 mm. for the total length of the specimen which is likely to have been the largest of the three. 7 v; x “ Yu wows WPS 7A. ALE aN Maas 4, Wie UR Te 5 CM. Fic. 17.—Tentoriceps cristatus (Klunzinger). Head of syntype ca 418 mm. S.L. (re-drawn after Klunzinger (1884), scale added). Some confusion is evident in the representation of the nostrils. Klunzinger’s figure shows only the head, pectoral fin and trunk back to the third dorsal ray. Even so the evidence available appears adequate to justify the recognition of a distinct species and genus, provided it is all accurately related and represented ; relatively small divergences from the published account would involve consideration of possible Assurger or Eugleurogrammus spp. Klunzinger’s careful consideration of the whole genus may justify confidence in his present data. It is obvious at the outset that T. cristatus has very little in common with the Aphanopodinae. It is likewise certain that, despite its ecaudate condition, it differs from the Trichiurinae in the general shape of the head, in the presence of ventral fins, in the median position of the lateral line and the absence of barbs from the teeth. (Elsewhere Klunzinger properly characterisises the barbed teeth and falling lateral line in Trichiurus muticus). 112 THE FAMILY TRICHIURIDAE Considered now as a possible Lepidopodine species 7. cristatus is quite unlike Lefidopus. The form of the upper profile of the head is intermediate between Evoxymetofton and Assurger, from both of which our species differs in lacking a caudal fin. From Klunzinger’s figure it seems that the elevation of the ethmo-frontal region has proceeded further than that of the posterior confluence of the frontal ridges giving an almost teratological appearance which is quite the reverse of the condition in the ecaudate Eupleurogrammus. The hind end of the operculum is a rounded point with an extension in relation to the pectoral base intermediate between that in the Aphanopodinae and Trichiurinae and unlike the other Lepidopodines. The number of dorsal rays (ca. 120) is similar to that in Assurger, but the elongation of the body, though considerable, is slightly less (depth 20-24: 28 in length). The number of dorsal spines is unknown, likewise the position of the ventral fin-insertions, the condition of the post-anal structures and the number of vertebrae ; nevertheless I find it possible to accept Tentoriceps cristatus (Klunzinger) as a valid species and genus arising from a Lepidopodine offshoot a little before Assurger. Whitley (1948) proposes Tentoriceps with no more than a translation of Klunzinger’s original description of Tvichiwrus cristatus, without any indication of the supposed discriminant characters and with no reference to his own earlier proposal of Assurger. He proposes it in a portmanteau paper of “ Studies in Ichthyology ” having no direct concern with the Red Sea fauna, no special interest in the Trichiuridae nor any Australian material of that family requiring comment. Tentoriceps is but another of Mr. Whitley’s foundlings, casually discovered, capriciously re-baptized and callously abandoned, in the hope of adoption or decent interment, on the cold doorsteps of systematic ichthyology. Subfamily TRICHIURINAE Swainson Trachiuvinae (evident misprint for Tvichiurinae) Swainson, 1839, Nat. Hist. Fish. Amphib. Rept. 2 : 254. Type genus Trichiurus Linnaeus. Lepturinae Gill, 1863, Proc. Acad. nat. Sci. Philad. 1863 : 225. Type genus Lepturus Artedi (= Trichiurus L.). GENERA NOW RECOGNISED: Tvichiurus Linnaeus ; Lefturacanthus Fowler. DIAGNOSIS : A. Slope of snout moderate ; orbits barely entering upper profile of head ; posterior confluence of frontal ridges elevated as a sagittal crest at the nape. B. Cartilaginous protuberance at mandibular symphysis weak; a small, soft projection at the tip of the snout. Lower hind margin of operculum more or less concave. Teeth of main series with barbs. Palatine teeth minute, in a villiform band. Lateral line descending steeply from the shoulder and running nearer the ventral surface of the body, i.e. distance between lateral line and ventral profile at anus slightly less than half distance between lateral line and dorsal, moO 4: THE FAMILY TRICHIURIDAE 113 G. Spinous dorsal fin very short, with 3 or 4 rays. Spinous and soft dorsals continuous, without any intervening notch. H. Soft dorsal rays precisely corresponding with adjacent caudal vertebrae, each basal and interneural element being related to a neural spine. I. Spinous anal i+ I; soft anal rays reduced to internal rudiments or wanting (Trichiurus) or taking the form of minute pungent spines which definitely break the ventral profile (Lepturacanthus). J. Anal fin (i.e., basal elements of anal—see I above) extending well beyond dorsal. K. Caudal fin and hypurals entirely absent. L. Ventral fins and girdle entirely absent. M. Pyloric caeca 24, perhaps more. Osteological literature Giinther, 1860, Cat. Fish. B.M. 2 : 343-344. (desc. osteology of Trichiurus). Starks, 1911, Stanford Univ. Pubs. No. 5 : 25-26 (desc. general osteology of Trichiurus, comp. with Lepidopus). Gregory, 1933, Tvans. Amer. Phil. Soc. 23 : 316, fig. 195 (skull of Trichiurus). Literature on young stages Delsman, 1927, Treubia 9 : 338. Liitken, 1880, K. Dansk. Selsk. Skrift. 12 : 409. Nair, 1952, Proc. Indian. Acad. Sci. 35B : 225. Tang & Wu, 1936, Lingnan Sci. J. 15: 651. Genus TRICHIURUS Linnaeus Trichiurus Linnaeus, 1758, Syst. Nat. Ed. 10 : 246. Type species Trichiurus lepturus Linnaeus ex Artedi (see note under T. lepturus). Pro- bably monotypic. Enchelyopus Klein, 1744, Hist. Piscium : 51. Enchelyopus Bleeker, 1862, Versl. Akad. Amsterdam 14 : 109. Type species Clupea haumela Forskal. (Also spelt Encheliopus by authors. Non Enchelyopus Gronovius, 1763). Gymnogastey Gronovius, 1754, Mus. Ichth. 1: 17. Type species Anguilla Jamaicensis Sloane. Lepturus Artedi, 1738, Desc. Spec. Pisc. : 111. Type species Lepturus argenteus Artedi. Lepturus Gill, 1863, Proc. Acad. nat. Sci. Philad. 1863 : 225. (Non Lepturus Moehring, 1758; Brisson, 1760.) ? Diepinotus Rafinesque, 1815, Analyse Nat.: 91 (nom. nud.). (Also spelt Dipinotus by authors.) ? Symphocles Rafinesque, 1815, Analyse Nat. : 91 (nom. nud.). ? Nemochirus Rafinesque, 1815, Analyse Nat. : 91 (nom. nud.). DIAGNOSIS : (1) Body-proportions highly variable : Head 7-0-9-4 in length, depth 14-4-21-0 in length. (2) Eye relatively large, 5-o-7-0 in head, 114 THE FAMILY TRICHIURIDAE (3) Dorsal spines III ; D.III, 137 in three specimens of three nominal species radiographed. (Published aggregate ranges D.120-140). (4) A.i+ I + 105-108. (5) Post-anal scute (= anal spine I) not enlarged; a small, triangular scale, less than the pupil. (6) First basal element of anal fin slightly enlarged, presumably a compound of 2, its interhaemal spine lengthened and slightly thickened. There follows a gap of I in the series of interhaemal spines, leaving 1 free haemal arch. (7) “‘ Soft anal’’ elements minute spinules which usually do not break the skin and which are occasionally absent. The first ca. 60 are directed backwards, the last ca. 40 are directed forwards. (8) Vertebrae 39-40 + 123-128 — 162-168. Probably only one variable species, Trichiurus lepturus L., world-wide except in colder regions. Trichiurus lepturus Linnaeus (Text-fig. 18). Trichiurus leptuyus (part) Linnaeus (ex Artedi), 1758, Syst. Nat. Ed. 10: 246. Type in the Museum of the Royal University of Upsala. Type locality South Carolina, Note.—Lénnberg et al., 1896, K. Svensk. Vet.-Akad. Handl. 22: 40, state that the Linnaean types of “ T. lepturus '’ at Upsala include material of the species now known as Eupleurogrammus muticus (Gray). The suggestion that T. lepturyus should consequently be replaced by T. argenteus Linnaeus, 1754, Mus. Ad. Frid. : 76, pl. 26, fig. 2 is, of course, illegal, nor is it really necessary since the situation has never created any practical difficulty. Giinther, 1898, Proc. Linn. Soc. Lond. 1898-9 : 20, satisfied himself that the Linnaean material in the possession of the Linnaean Society of London is, in fact, T. leptuvus, which is rendered doubly certain by the fact that it came from Garden’s South Carolina collections, consignment of 1761. (See also id. ib. : 25.) Trichiurus lepturus J. L. B. Smith, 1949, Sea Fish South Africa : 313; Okada, 1955, Fishes of Japan : 155. Trichiurus argenteus Shaw, 1803, Gen. Zool. 4: go, pl. 12 (apparently ex Linnaeus, 1754). Clupea haumela Forskal, 1775, Descr. Anim. : 72. Type not in Herbarium Ichthylogicum Forskalii, Copenhagen (fide N. B. Marshall, personal communication). Type locality Red Sea. Trichiurus hamrela Schneider, 1801, Syst. Ichth. : 518 (nom. err.). Trichiurus lepturus japonicus Temminck & Schlegel, 1844, Faun. Jap. Pisc. : 102, pl. 54. Type Leiden Museum No. 2040. Type locality Japan. Trichiurus lepturus japonicus Boeseman, 1947, Zool. Meded. 28: 96 (for Temminck co-author p: 2). Trichiurus japonicus Bleeker, 1857, Verh. Bat. Gen. 26: 08. Trichiurus japonicus Lin, 1936, Bull. Chekiang Fish. Sta. 2 (5) : 2. Trichiurus lajoy Bleeker, 1854, Nat. Tijdschr. Ned. Indie 7 : 248. Type in Leiden Museum. Type locality Manado, Celebes. (Re-examined by De Beaufort, 1951, Fish. Indo-Austr. Archip. 9 : 196.) Trichiurus malabaricus Day, 1865, Proc. zool. Soc. Lond. 1865 : 20. Holotype B.M. (N.H.) No. 1867.5.30.2. Type locality Madras. (Withdrawn as T, haumela by Day, 1876, Fish, India : 201.) FAMILY TRICHIURIDAE THE i 116 THE FAMILY TRICHIURIDAE Trichiuvus auviga Klunzinger, 1884, Fische Rothen Meeres 1: 120, Pl. 12, fig. 1. Type retained in Klunzinger’s private collection, Stuttgart. Now at Stuttgart, Berlin or Vienna? Type locality Kosseir, Red Sea coast of Egypt. Trichiurus auriga Weber, 1913, Siboga Fische : 406. Trichiurus auriga De Beaufort, 1951, op. cit. : 196. Trichiurus coxit Ramsay & Ogilby, 1887, Proc. Linn. Soc. N.S.W. 1887 2 (2) : 562. Holotype Australian Museum, Sydney No. 1.1342. Type locality Broken Bay, N.S.W. ? Trichiurus nitens Garman, 1899, Mem. Mus. comp. Zool. Harv. 26 : 69. Syntypes (2) in U.S. N.M.? Type locality coast of Peru. ? Trichiurus nitens Hubbs & Hubbs, 1941, Calif. Fish Game 27 : 29. ? Trichiurus nitens Breder, 1936, Bull. Bingham Ocean. Coll. 2 Art. 3 : 12. non Trichiurus lepturus Mohr, 1786, Forsog til en islandsk Naturh. Kjob. : 63. non Trichiurus lepturus Sveinn Palsson, 179+, J. Naturforsk. Reise Island 1791-97 2. (Mis-identifications of Tvachypterus sp. Refs. fide Saemundsson, 1926, Fiskaynir : 155. Reykjavik). non Trichiurus lepturus Hoy, 1815, Trans. Linn. Soc. Lond. 11 ; 210-212. (Mis-identification of Tvachypterus or Regalecus spp.) non Trichiurus tvimaculatus Giovene, 1829, Mém. Soc. Ital. 20 Pt. 1: 25. (Mis-identification of Tvachypterus sp.) At the commencement of the present paper Trichiurus seemed likely to give the most difficulty ; that promise has been abundantly fulfilled. Tvichiwrus is a common pelagic fish of world-wide distribution, occurring in all but the coldest seas and assuming some economic importance in certain areas ; as a consequence it possesses a literature as large as that of the rest of the family put together. Much of this work is uncritical : species have been recognized on supposed differences of body-proportions unrelated to possible ontogenetic changes, geographic variation, or environmental effects, or on small differences in fin-ray counts which are difficult to establish with any accuracy except in radiographs. Vertebral counts have hardly ever been employed. Very often it is found that where a worker has characterised a pair of species to his own satisfaction another will reverse the discriminant characters in the same pair. In dealing with all this intractable material it has seemed useful to take as a working hypothesis the theory that we are dealing with one highly variable species. If the evolutionary behaviour of the other recent Trichiuridae affords any precedent it is one pointing to the evolution of monotypic genera, or of pairs of species having sharply discontinuous ranges of meristic counts, not to the subtle distinctions which the would-be splitters of Tvichiurus postulate. Geographic variation and theincreasing evidence of environmental effects upon meristic characters must also be taken into account. The first problem is the identity or distinctness of the Atlantic and Indo-Pacific populations; currently recognized as 7. lepturus L. and T. haumela (Forskal) respectively. The results of the examination of two specimens taken at random from the collections of the British Museum (Natural History) are given in Table V. They show a precisely coincident dorsal count and anal/vertebral counts differing by only 3/2 rays/vertebrae respectively. The differences in body proportions are no greater than may be explained by the difference in age. These two specimens show that similar Tvichiurus occur off Texas and Shanghai almost as well as population samples treated with all the apparatus of statistical necromancy. THE FAMILY TRICHIURIDAE 117 TABLE V. Trichiurus Trichiurus Trichiurus lepturus. “ haumela.” “japonicus.” Aransas Bay, Texas. Shanghai. Yenting, Chekiang. No. 1948.8.6.795. No. 1862.11.1.260. No. 1925.4.23.5- Dorsal rays c : D.III, 137 a D.III, 137 2 D.III, 137 Analrays . : 3 A.i+I+105 c Avi+I-+108 0 A.i+I+107 Vertebrae . : 2 39 +123=162 : 40 +124=164 c 40 +128=168 Standard length . c 545 mm. c 838 mm. : 926 mm. Head in S.L. : é 7:07 7 7°83 c 9°17 Depth in S.L. : : 16°03 a 16°43 é 17°47 Snout-vent in S.L. : 2-79 2 3°07 5 Bor) Head in S-V. 5 . 2°53 5 2°55 2°98 Depth in S-V. 6 : 5°73 5°35 5°68 Eye in head 5°92 6-68 6:12 Snout in head 2°80 iy) 2°52 Mx. in head 2°48 2°46 2°29 A modern re-assessment of Trvichiurus japonicus Temminck & Schlegel, a second Pacific form, is given by Boeseman (1947) : “ The differences between Trichiurus japonicus T. & S. and T. lepturus L. as stated by Temminck & Schlegel and Bleeker (/.c.) do not exist in our material. A comparison, however, with several specimens of /eptwrus in our collection (all Atlantic) showed a very distinct and constant difference ; the head in all these specimens of lepturus is larger, about 7-7-5 in length, while in our specimens of japonicus it is 8-1-8-6 in length, consequently considerably smaller. On account of this I want to discriminate both species and regard the Japanese specimens as the type material of a separate species, Trichiurus japonicus T. & S. Specimen no. 2040 I regard as type.” In opposition to this stands the work of Lin (1936) who comments that : “Several authors ... used to distinguish T. japonicus from T. hawmela by the shorter head and the smaller eyes, but the series of intermediate forms lying between them is so continuous that no clear line can be drawn to separate them into two distinct species.” Lin gives a table of data covering a series of 12 Chinese T. japonicus of 424-1,290 mm. S.L., with the following ranges : depth in length 14-4~21 ; head in length 7-9-4 ; depth in head 1-9-2-3 ; eye in head 5-4-6-4; D.136-140. He notes the difficulties which arise through the loss of the tip of the tail and advocates the substitution of snout-vent length for the calculation of body-proportions. Oddly enough Lin does not take the logical step of substituting the earlier name hawmela for japonicus. “In reviewing the recorded distribution of T. jafonicus and T. haumela, it is found that the former species inhabits the Chinese and Japanese coastal waters and is neither known to live beyond the Asiatic continental shelf nor 118 THE FAMILY TRICHIURIDAE southward to the East Indian seas, while the latter is found from the Japanese and Chinese seas to the Philippines and Indian Ocean and Archipelago.” A specimen of T. japonicus (Table V), being duly radiographed, shows a dorsal count identical with that of the two specimens of the other two nominal species, an intermediate anal count and an increment in caudal vertebrae altogether less than the head/body proportion would lead one to expect. I suggest that T. japonicus is, in the light of the evidence, no more than an ecotypic form of T. hawmela (= lefturus). T. lajor Bleeker and T. malabaricus Day have been adequately dealt with by De Beaufort (1951) and by Day himself (1876) and as synonyms of 7. haumela now need no further comment. T. auriga Klunzinger, placed very close to T. haumela by Klunzinger (1884) himself and founded on a very young specimen (250 mm. S.L.) is probably no more than a juvenile of the latter species, though the published illlustration contains peculiar features at variance with the description. The only serious difficulty is the definitely stated absence of barbs from the teeth. 7. awriga has been reported only once more, a specimen of 320 mm. from the Timor Sea having been described by Weber (1913) and De Beaufort (1951). T. coxtt Ramsay & Ogilby (1887) contains, on the published account, no differences from 7. hauwmela and its authors do not attempt to indicate those characters in which they consider it to be divergent. Only in the case of 7. nitens Garmen (1899) has there been argued any very cogent case for specific separation, by Breder (1936) and by Hubbs & Hubbs (1941). Breder’s conclusions, summarized in the form of a key are: A. Dorsal rays never less than 126, usually about 133 ; maxillary 2'2 to 2°5 in head, usually about 2° 3 (11 specimens, D.126-137, mean 132*9) . : . lepturus Atlantic AA. Dorsal rays never more than 128, usually about 122; maxillary 2'5 to 2°8in head, usually about 2°6 (36+ specimens, D.120-128, mean ca. 122°3 nitens California, Galapagos These results must obviously be treated with respect, though not with absolute aquiescence. The variation in the maxillary should be stated in relation to the size of the fish, for there is some allometry with age. The differences in mean dorsal counts are considerable, but if. the two samples are drawn from a continuous hypo- thetical population of one species, having its origin somewhere in the Indian Ocean and extending westwards to the Western Atlantic and eastwards to the Pacific coast of America we still have no more than the variation we should expect at the limits of that wide range. The satisfactory establishment of T. nztens requires, not compari- son with Atlantic material, but demonstration of a non-cline discontinuity across the Pacific. Having regard therefore to the known variation in the Trichiuridae and to the growing literature concerning environmental effects upon fishes, I am more inclined to “Jump ”’ Tvichiuwrus as one variable species than to “ split’ and thereby make assumptions concerning the genetic distinctness of populations which present precedents and evidences of variation do not appear to justify. Such modern authors as J. L. B. Smith (1949) and Okada (1955) have apparently arrived at the same conclusion, though they do not submit any evidence to support their decisions. THE FAMILY TRICHIURIDAE 119 Genus LEPTURACANTAUS Fowler Lepturacanthus (sub-genus of Trichiurus L.) Fowler, 1905, Proc. Acad. nat. Sci. Philad. 1904 : 770. Type species Trichiurus savala Cuvier. Monotypic. Trichiurus (part) many earlier authors. DIAGNOSIs : (t) Body-proportions highly variable: Head 7-4-10°5 in length, depth 14-8- 19°8 in length. (2) Eye relatively small, 6-7—10-0 in head. (3) Dorsal spines IV; D.IV, 111 in two specimens radiographed, including holotype of Tvichiurus armatus Gray. (Published aggregate ranges D.105-134). (4) Ai+ 1+ 72. (5) Post-anal scute (= anal spine I) enlarged, as in Aphanopus ; a dagger-like spike half the diameter of the eye. (6) First basal element of anal fin markedly enlarged, as in Aphanopus, presumably a compound of 3, its interhaemal spine likewise lengthened and thickened. There follows a gap of 2in the series of interhaemal spines, leaving 2 free haemal arches. (7) “Soft anal” elements pungent spinules, definitely breaking the ventral profile throughout the length of the fin and all directed backwards. (8) Vertebrae 32-35 + 124-130 = 159-162. One species, Lepturacanthus savala (Cuvier). Indo-Pacific. Lepturacanthus savala (Cuvier) (Text-fig. 19). Trichiurus savala Cuvier, 1829, Régne Animal 2 Ed. 2 : 219. Syntypes in Paris Museum, Reg. No. a.5357-5358. Type locality ‘‘ Mer des Indes ” (= Bombay & Malabar). Trichiurus (Lepturacanthus) savala Fowler, 1905, Proc. Acad. nat. Sci. Philad. 1904 : 770. Trichiurus armatus Gray, 1831, Zool Misc. 1:9; Gray, 1835, Illust. Ind. Zool., pl. 93, fig. 1. Holotype B.M. (N.H.) No. 1955.5.13.1. Type locality India. Trichiurus Roelandti Bleeker, 1860, Acta Soc. Indo-Neer. 8 (4) : 30. Holotype in Leiden Museum. Type locality Sunda Strait. (Re-examined by De Beaufort, 1951, Fish. Indo-Austr. Archip. 9 : 194.) Trichiurus armatus Gray has for long, and, for once, correctly, been regarded as a synonym of this species. De Beaufort has adequately dealt with T. Roelandti Bleeker and so the taxonomic situation in this newly-promoted genus Lepturacanthus is mercifully straightforward. Lepturacanthus is obviously closely related to Trichiurus and as widely separated from the other Trichiuridae : any attempt at a natural classification must adequately express this situation. In a wide classification Fowler’s erection of Leptwracanthus as a sub-genus of Trichiurus very adequately did so, but with the exposure of Eupleuro- gvammus and its removal to the Lepidopodinae the Trichiurinae are left as a very ZOOL, 4, 3- 9 120 THE FAMILY TRICHIURIDAE small group. Within this group the divergence between Lepturacanthus savala and Trichiurus lepturus is very comparable to that between A phanopus and Benthodesmus and accordingly consistency requires the elevation of Leptwracanthus to full generic status. THE ORIGIN, EVOLUTION AND CLASSIFICATION OF THE TRICHIURIDAE Summary of earlier work Before summarizing previous opinions on the classification of the Trichiuridae it may be useful to indicate the sequence of recognition of the genera now accepted : Trichiurus Linnaeus, 1758. Lepidopus Gouan, 1770. Aphanopus Lowe, 1839. Evoxymetopon (Poey) Gill, 1863. Eupleurogrammus Gill, 1863. Benthodesmus Goode & Bean, 1882. Lepturacanthus Fowler, 1905. Assurger Whitley, 1933. Diplospinus Maul, 1948. Tentoriceps, Whitley, 1948. Classification commences, and commences remarkably well, with Cuvier & Valenciennes (1831) who recognize the Scombroids as a natural group (Scombéroides) containing all the Trichiurids and Gempylids so far known (Lépidopes, Trichiures ; Thyrsites, Gempyles) as well as the Tunnies, etc. Their key runs : “ Tous ou une grande partie des rayons de l'anale véduits a de trés-petites épines. Dents des thyrsites et des gempyles. LePIpoPEs. Une petite écaille au lieu de chaque ventrale ; une caudale. TRICHIURES. Point de ventrales ; point de caudale.”’ These authors grasp so early the essential relationships of the Trichiuridae and Gempylidae : “Tl est impossible de ne pas placer 4 la suite des gempyles et des thyrsites deux genres de poissons qui leur ressemblent presque en toutes choses, si ce nest qu’ils manquent entiérement de fausses nageoires et méme de rayons mous A leur dorsale; ce sont les lépidopes et les trichiures, poissons trés- remarquables d’ailleurs par leur éclat et par leurs formes singuliéres. “Leur téte, leurs dents, leur peau, leur squelette, rappellent de tout point les genres auxquels nous les associons, et la longueur méme de leur corps en ruban, qui les avait fait rapprocher des cépoloides, est déja annoncée par la forme de plusieurs gempyles.”’ Swainson (1839) was the first to erect a higher taxon for the Trichiurid fishes, though he takes a step backwards, making Ammodytes as well as Trichiurus and Gempylus members of a sub-family Trichiurinae of the family Coryphaenidae. THE FAMILY TRICHIURIDAE 121 Giinther (1860) attempts no subdivision of his family Trichiuridae, which includes not only Aphanopus, Lepidopus and Trichiurus but also the Gempylids Epinnula, Dicrotus, Thyrsites and Gempylus. Gill (r863) gives a classification recognizably approaching that now advocated, though based on inadequate and in part inaccurate premises : “J. Dorsal fin undivided. A. Tail filiform and finless. : : 0 : . LEPTURINAE. Lateral line near the abdomen. ; : : A . Lepturus. Lateral line median 5 c - . Eupleurogrammus. B. Tail with a normally developed and forked fa c 0 . LEPIDOPODINAE. Profile rectilinear and forehead depressed : c . Lepidopus. Profile high, trenchant and mae. Sao 0 6 . Evoxymetopon. II. Dorsal fin double. “ : . APHANOPODINAE. Teeth of the palate wanting 5 ¢ c : c . Aphanopus. Johnson (1865) describes a number of Gempylids as Trichiuridae. Capello (1868) takes a view of the Trichiuridae equivalent to the Trichiuriformes of later authors and recognizes three sub-families : Trichiurus, Eupleurogrammus. TRICHIURINA : : ; : Lepidopus, Evoxmyetopon. Aphanopus. GEMPYLINA . : : : : Gempylus, Prometheus, Epinnula. THYRSITINA . : . : : Thyrsites, Dicrotus. A division of the Trichiurina similar to Gill’s is implicit in the key given. Time, on the whole, has dealt more kindly with Capello than did Giinther in the Zoological Record. Goode & Bean (1895) limit the Trichiuridae to Tvichiwrus, of which Ewpleurogrammus is merely ‘“‘a Chinese form ... with a single species”! They erect a separate family, the Lepidopidae, with two sub-families : “I. Dorsal continuous. Teeth on palatines. Ventrals present, scale-like, rudimentary. No post-anal spine 3 4 : LEPIDOPINAE. (Genera Lepidopus, Bete Feninadere Il. Dorsal in two subequal portions, closely contiguous. No teeth on palatines. Ventrals absent. A dagger-like post-anal spine = 5 6 < APHANOPINAE. (Genus Aphanopus.) This pastiche of half-truth and etymological abomination is preceded by one of Goode & Bean’s self-contradictions (Lepidopidae have “ No teeth on palatines ”’ Lepidopinae have “ Teeth on palatines ’’). Boulenger (1904) and Goodrich (1909) include both Trichiurids and Gempylids in a family Trichiuridae without subdivisions. This grouping becomes the Scombroid Division Trichiuriformes of Regan (1909), with two undivided families Trichiuridae and Gempylidae in the generally accepted modern sense, as later followed by Jordan (1923) and by Berg (1940). Starks (1911), in a classic paper concerned only with the osteology of three genera, defines families Gempylidae (Promethichthys), Lepidopidae (Lepidopus) and Trichi- ZOOL. 4, 3. 9§ 122 THE FAMILY TRICHIURIDAE uridae (Trichiwrus). He comments that ‘‘the descent of the family Trichiuridae from the Gempylidae was long ago pointed out’ and compares the structure of Lepidopus with that of Promethichthys, which latter he rightly regards as too specialized to be an ancestral form. He concludes: “‘ This ancestor may have been Gempylus, a form which I have been unable to obtain, but showing a development towards the elongate forms of Lepidopus and Trichiurus.” Roule (1927) introduces a little light relief by attempting to place the Iniomous Anotopterus among the Trichiuridae. Gregory (1933) figures a museum exhibit showing in pictorial form the evolution of the Scombroid fishes. Ruvettus, Epinnula, Gempylus and Trichiurus are shown as consecutive stages in a linear series, with Lepidopus emerging as a sideshoot between Epinnula and Gempylus. In these circumstances the presence of a number of apparently undecided fishes swimming in the background to this exhibit occasions no surprise. Tucker (1953), though not attempting a full classification, draws attention to the affinities between Aphanopus and Benthodesmus as contrasted with Lepidopus. He shows that Benthodesmus has a differentiated and partly divided dorsal fin like that of Aphanopus and demonstrates the significance of the ventral fin-insertions and post-anal structures, but, fails to realize that the dorsal fin is differentiated throughout the entire family. The error arose through an undue reliance on previous literature of the non-Aphanopodinae and a brief study of Lepidopus and Trichiurus from radiographs which, for reasons of economy, were fragmentary. Dr. Carl L. Hubbs, in litt. kindly drew attention to this mistake. Nesiarchus-Diplospinus : the Gempylid-Trichiurid bridge Regan (1909) gives the following diagnosis of the Gempylidae which may still serve as a basis for comparison with the Trichiuridae (p. 74) : “Body oblong or elongate, compressed ; maxillary exposed ; spinous dorsal longer than the soft ; anal with 3 spines, similar to the soft dorsal ; each pelvic fin of a spine and 5 soft rays or reduced to a spine only ; caudal fin present. Rays of the spinous dorsal equal in number to the vertebrae below them, each interneural usually attached to a neural spine; rays of soft dorsal and anal more crowded (except the isolated finlets, when present), about twice as numerous as the corresponding vertebrae ; pelvic bones separate, anteriorly extending forward to the cleithra and firmly imbedded in the ligament between them. Vertebrae 31(15 + 16) to 53(28 + 25); anterior praecaudals without parapo- physes, with sessile ribs; posterior praecaudals with ribs attached at the extremities of closed haemal arches ; epipleurals attached to the centra.” Closely related to the Scombridae, from which, however, they may be descended by more than one line, the Gempylidae are quite a varied group of fishes. As Mrs. Grey (1953) notes : “There is a puzzling scattering of such characters as the presence of a free dagger-shaped spine preceding the anal fin, of dorsal and anal finlets, double or single lateral lines ; and the presence, absence, or reduction of ventral fins.” THE FAMILY TRICHIURIDAE 123 A goodly proportion of the genera are well-illustrated in the paper by Matsubara & Iwai (1952). The ancestors of the Trichiuridae must undoubtedly be sought among the Gempy- linae (Gempylus Cuvier, Nesiarchus Johnson, Mimasea Kamohara), Gempylidae which possess an especially elongate body, the head and trunk in particular being reminiscent of those of the Trichiurids although the tail seems greatly telescoped by comparison and curiously unfinished. In these three genera alone appear the conical cartilaginous processes at the tip of the snout and mandibular symphysis which are found in the Aphanopodinae; their skulls are long and low, without prominent crests; they have, like other Gempylids, the typical Trichiurid dentition with the three pairs of prominent premaxillary fangs ; their squamation is, at the most, vestigial, leading directly to the naked bodies of the Trichiuridae. A single row of teeth is present on the palatine in Nesiarchus (personal observation) and in Gempylus (Matsubara & Iwai), though Mimasea is said to have none. Of these three Gempyline genera Mimasea (Text-fig. 20) is specialized in having a double lateral line and a ventral fin-insertion behind the pectoral base ; primitive in that the ventral fin is quite well developed, with five soft rays. Despite low median fin-ray counts and presumably low vertebral counts therefore, it does not seem a likely ancestor to the Aphanopodine Trichiurids. Gempylus (Text-fig. 21) has rather less well-developed ventrals, allied, however, to a double lateral line and a series of widely-spaced dorsal and anal finlets. The proportion of soft dorsal rays to aggregate vertebrae in this genus is very low (18 : 53) in comparison with Diplospinus, the most primitive recent Trichiurid (40 : 58) and, since the early history of the Trichiurids appears to show soft dorsal rays multiplying much faster than the caudal vertebrae, we may feel that the transition from 53 to 58 vertebrae represents a smaller change than is likely to admit the necesary concomitant structural changes (Table VI). Nesiarchus, (Text-fig. 22) however, seems to stand very close to the primitive Trichiuridae. It has a total of vertebrae (35) near to the minimum of its family (3r in Epinnula), allied to a higher number of soft dorsal rays than in Gempylus (21-23 : 18) and without detached finlets. The ventral fins are inserted on the per- pendicular through the posterior end of the pectoral base and consist each of a spine with four smaller soft rays. The skull is well figured by Steindachner (1867) ; apart from a broad general resemblance to the skulls of the Aphanopodinae there is a striking similarity in the deep opercular notch, nowhere as marked in the other Gempylidae and found in only one Trichiurid—the primitive Diplospinus. The post-anal spines appear superficially “wrong” ; the first is much larger. But internally there is a rudiment of yet another before the first ; there are three well-developed spines in Epinnula and it becomes evident that of these the first is to become the minute spinule of Nesiarchus and the Trichiuridae (i), the second will become the larger spine of Nesiarchus and the principal spine or scute of the Trichiuridae (I), and the third, though disappearing, is to signify its claim by a space in the anal fin and will contribute to the compound and reinforced anterior basal structure whenever this is developed, — = “QIIM ‘UT O00'T “ MWS obo ON FS LT“ AOBTI 4S Wo _ euRg ,, Woy “T'S ‘WU zgr jo ueuvads “uosuyof snynsvw snysavisaN—ZE “OTT *(zS6r ‘remy 2% eIeQns}eW WOIJ) oLUeS Jo UY [eIZUEA fo [IeJaq “A ‘uede{ “J's ‘uu ggh yo usumoeds “vy “JoIAND suadaas snjpkdmag— iz “91d THE FAMILY TRICHIURIDAE *(zS6r ‘remy 2 eIeGns}ey[ WOI,) uedef “J's WU ogz ‘eIeYOUIRy YULOSOIUaD, VaSDULITY—'O% “OTT 124 THE FAMILY TRICHIURIDAE 125 The body of Nesiarchus is quite naked ; the lateral line is single and descends gently to a mid-lateral course along the caudal. The number of pyloric caeca (7) is similar to that in the Aphanopodinae. Nesiarchus differs from Diplospinus in the lower meristic counts, in having the maxillary exposed, in having barbs on the teeth confined to the premaxillary fangs and in the external (though not in the internal) structure of the spinous anal fin. But the indications from a study of the subsequent evolution of the Trichiuridae are that during the addition of 18 vertebrae Nesiarchus would have had plenty of time to undergo the modifications needed to produce a Diplospinus. This is the view expressed in Text-fig. 23. Nesiarchus and Diplospinus therefore may be regarded as the approaches to the Gempylid-Trichiurid bridge. Whether the Trichiurinae crossed by the same bridge or by a parallel bridge further downstream is still debatable. It is tempting to regard the low lateral line in the Trichiurinae as representing the lower limb of the fork in another Gempylid ancestor; but unfortunately, although Trichiurus has a longitudinal groove which would serve for an upper limb, no recent Gempylid has a lower limb which falls in quite the same way. If, however, the “ toothless ’’ palatines in Mimasea should, on further examination, prove to be provided with a villiform band of teeth, the discovery would be a significant indication of a possible relationship and therefore of a diphyletic descent of the Trichiuridae. In this connexion it is interesting to observe that the concave lower hind margin of the operculum, charac- teristic of the Trichiurinae (though not of Nesiarchus, the Aphanopodinae or the Lepidopodinae), makes sporadic appearances among the primitive Gempylidae in Epinnula and Neoepinnula. Evolutionary trends in the Trichiuridae Evolution in the Trichiuridae has resulted from the action, at various rates, of the following trends : (x) Elongation of the caudal region of the body, least in the stem-forms at any level (Diplospinus, Lepidopus) and greatest in the most divergent side-shoots (Benthodesmus, Assurger, Tentoriceps). (2) Multiplication of the soft dorsal and anal rays, initially at a greater rate than that of the adjacent vertebrae. This development, already incipient throughout the Gempylidae, is seen proceeding at its greatest rate in Diplospinus and is practically in harmony with the multiplying vertebrae in the other Aphanopodinae. (3) Multiplication of the caudal vertebrae until eventually (except at the caudal tip) each vertebra has one corresponding soft dorsal and anal ray with their associated basal elements. This process is nearly complete in Aphanopus and Benthodesmus, in which, however, there are usually a very few rays, mainly towards the beginning and end of the soft fins, which are not directly related to vertebrae. Except possibly in “ Lepidopus xantusi”’ further additions of vertebrae and fin rays proceed in unison in the Lepidopodinae and Trichiurinae. (4) Aslower increase in the number of trunk vertebrae. (See Table VI in conjunction with Text-fig. 23). 126 THE FAMILY TRICHIURIDAE LEPIDOPODINA Meter Trichiurus Lepturmcanithus Benthodesmus TRICHIURINE APHANOPODINE — Diplospinus Nesiarchus GEMPYLINA. Fic. 23.—Suggested relationships of the genera of the Gempylid subfamily Gempylinae and of the subfamilies and genera of the family Trichiuridae. THE FAMILY TRICHIURIDAE 127 TABLE VI. Dorsal. oO Vertebrae. Spines. Soft rays. GEMPYLIDAE Gempylinae : Nesiarchus nasutus 0 19-21 21-23 : 23 +12=35 Mimasea taeniosoma . - 18 16-18 0 — Gempylus serpens cS : 29-32 18 A 53 TRICHIURIDAE Aphanopodinae : Diplospinus multistriatus 32-33 40 . 34 +24=58 Aphanopus carbo . : 0 38-41 53-50 a 42-44 +55-56=98-99 Benthodesmus tenuis. c 39-42 80-88 7 47-52 +75-80= 123-131 Benthodesmus simonyi . 4 45-46 102-108 + 52-53 + 101-103 =153-156 Lepidopodinae : “Lepidopus xantust”’ . ; (82) ¢ — Lepidopus caudatus 6 0 9 90-97 c 41 +70-73=I1I-113 Assurgey anzac . c A (ca. 120) 2 _ Tentoriceps cristatus . : (ca. 120) c — Evoxymetopon taeniatus : 10 77 c — Eupleurogrammus intermedius 3 123-131 - 32-35 +125-128=157-162 Eupleurogrammus muticus . 3 143-147 : 41 +150-15I=IgI-192 Trichiurinae : Trichiurus lepturus 6 3 137 - 39-40 +123-128= 162-168 Leptuvacanthus savala . 6 4 III + 32-35 +124-130=159-162 Data for Nesiarchus (part), Mimasea & Gempylus taken from Matsubara & Iwai (1952) and Grey (1953) ; for ‘‘Lepidopus xantusi” from Jordan & McGregor (1899) ; for Assurgey anzac from Alexander (1916) ; for Tentoviceps cristatus from Klunzinger (1884) ; for Evoxymetopon taeniatus from Gill (1863). Remainder original. (5) Progressive reduction of the number of dorsal spines in the higher forms and their replacement by soft rays. (6) Backward migration of the ventral fins. In the Gempylidae the pelvic girdle is embedded in the ligament between the cleithra, and the primitive position of the ventral fin-insertions in the Trichiuridae is likewise closely before or behind the level of the pectoral base. In Benthodesmus simonyi they are already further back than in B. tenuis, while among the Lepidopodinae the migration continues : an eye-diameter behind the pectoral base in Lepidopus and Evoxymetopon, an eye-diameter and a half in Assurger and finally five eye-diameters in Eupleurogrammus, but all the while a ligamentous connection between the pelvic girdle-rudiment and the symphysis is maintained. This situation has been discussed by Regan (1909 : 67). (7) Hypertrophy of the dorsal musculature, with consequent elevation of the posterior confluence of the frontal ridges of the skull into a distinct sagittal crest (Lepidopus, Trichiurus) followed at a later stage by an adjacent elevation of the ethmo-frontal region continuing this crest forward along the snout (Evoxymetopon, Assurger, Eupleurogrammus, Tentoriceps). (8) Increase in the number of pyloric caeca. In the Aphanopodinae, as in the 128 THE FAMILY TRICHIURIDAE Gempylinae, the number lies within the range 6-9 (6-8 in 30 Aphanopus counted) ; in Lepidopus it is over 20 and may be much higher in Tvichiwrus (which needs to be studied from fresh material). (9) Reduction of the soft anal fin from before backwards. In Diplospinus the soft anal extends nearly to the vent, so also in Benthodesmus tenuis. In Aphanopus and in B. simonyi the anterior rays are weak and probably of no functional consequence in the fin ; in Lepidopus and others only the last 20 rays or so form the true fin. In Lepturacanthus and in Trichiurus the whole fin is reduced to a series of minute spinules, while in Ewplewrogrammus the fin as such has ceased to exist and only the basal and interhaemal elements remain, firmly interlocking with the haemal arches to form a continuous mid-ventral keel. (x0) Loss of the caudal fin and hypural bones, independently in Trichiurus, Lepturacanthus, Eupleurogrammus and Tentoriceps. (11) Reduction in the extent of the intermuscular (pleural and epipleural) bones. In Diflosfinus these long bones are a prominent feature in the skeleton and form a complete basket surrounding the abdominal cavity, but in all the other genera the space which they contain becomes a much smaller portion of the whole. In Eupleuro- gvammus a small “‘ basket ’’ supported by 14 rather smaller vertebrae is pushed to the anterior end of the trunk and is followed by 18-25 vertebrae without epipleurals. Pari-passu with the major trends outlined above come sporadic tendencies, repeated at different levels : (a) Excessive elongation of the body, a possible symptom of evolutionary inertia (Benthodesmus, Assurger, Tentoriceps). (b) Hypertrophy of the second anal spine, with correlated condensation of the anterior basal and interhaemal elements into an enlarged supporting structure. (Aphanopus, Lepturacanthus). (c) Reduction of the pelvic girdle and fins to an internal rudiment (A phanopus) or their complete loss (Tvichiurus, Lepturacanthus). Classification of the Trichiuridae The Aphanopodinae as now recognised comprise Gill’s group (Aphanopus) with the addition of Benthodesmus and Diplospinus, genera recognized since Gill’s time. They are forms in which the major changes from the Gempyline condition have been accomplished but in which the evolution of the Trichiurid caudal may still be seen proceeding. The discriminant characters of the primitive Diplospinus have already been noted ; it isa satisfactory ancestral form except possibly in the advanced barbing of the teeth, a character which, if not merely adaptive, may indicate an affinity with the ancestors of the Trichiurinae rather than with the Nesiarchus- Aphanopus line. Aphanopus is a secondarily specialized bathypelagic form having an enlarged postanal spine and associated endoskeleton. Benthodesmus is an attenuate type which has gone some way with Aphanopus (as evidenced by the endoskeleton of the anterior anal fin) and then stopped. B. simonyi, evidently derived from B. tenuis, shows several evolutionary trends in action in the same genus. The Lepidopodinae are equivalent again to Gill’s group (Lepidopus, Evoxymetopon) with the addition of Eupleurogrammus (removed from Gill’s Lepturinae = THE FAMILY TRICHIURIDAE 129 Trichiurinae) and of other genera subsequently recognised—A ssurger, Tentoriceps, Lepidopus (as represented by L. caudatus) shows a great reduction in the spinous dorsal and the early stages in the uplift of the cranial crest and in the backward progress of the ventral fins; at the same time it has attained equilibrium in the development of vertebrae and soft fin-rays, and is well on the way towards losing its anal fin. The so-called “‘ Lepidopus xantusi’’ of unhappy memory is inadequately known, but would appear to be more primitive than L. caudatus and may even deserve generic status in a position between Lepidopus and Diplospinus in the main stem. In my opinion L. caudatus represents the termination of a very old line and its close similarity of skull to Trichiwrus is the result of parallelism and not of any closer relationship. The remaining Lepidopodine genera—Evoxymetopon, Assurger, Tentoriceps, Eupleurogrammus—have in common an elevation of the ethmo-frontal region to continue the sagittal crest forward from the nape to the snout ; in Eupleuro- grammus, the only one of this quartet which I have been able to handle, the homo- logies in relation to Lepidopus are easily discernible and, together with published figures, give sufficient indication of the likely condition in the other three. Evoxy- metopon is probably the most primitive of this group, in its shorter body and lower median fin-ray counts and in the position of the ventrals and presence of a caudal fin, but has a rather steep profile. Theecaudate andhighly perfected Eupleurogrammus may have been descended from this line, sharing with the Lepidopodines (and not with Trichiurus, with which it was formerly classified) the uniseriate palatine teeth, median lateral line, ethmo-frontal elevation, ventral fins and rounded operculum. The elongate, caudate Assurger and the ecaudate Tentoriceps form another like pair. The Trichiurinae are now restricted to Trichiurus and Lepturacanthus, the latter Fowler’s sub-genus upgraded to full generic rank. They are unique among the Trichiuridae, not for their loss of a tail (which has occurred elsewhere and indepen- dently), but in having a band of villiform teeth on each palatine rather than a single series, in having lost the last vestige of a pelvic girdle and fins and in having a low- descending lateral line. Other differences assume greater significance in relation to these. It is therefore likely that the fundamental cleavage between the Trichiurinae and the other two sub-families goes deeper than has previously been supposed. It is interesting to observe, in conclusion, that although there has been such a great reduction in the number of nominal species formerly placed in Tvichiurus the residue are now distributed through five genera—Lepidopus, Trichiurus, Leptura- canthus, Eupleurogrammus and Tentoriceps. REFERENCES (Excluding those given under synonymies of genera and species.) Bere, L. S. 1940. Classification of fishes, both recent and fossil. Tvav. Inst. Zool. Acad. Sci. U.R.S.S. 5 : 87-517, figs. Boutencer, G. A. 1904. Teleostei. Cambridge Natural History 7 : 539-760, figs. London. CAPELLO, F. DE B. 1868. Catalogo dos Peixes de Portugal que existem no Museu de Lisboa. J. Sci. Math. Phys. Lisboa 1 : 233-264, pl. 130 THE FAMILY TRICHIURIDAE Cuvier, G. & VALENCIENNES, A. 1831. Histoive Naturelle des Poissons 8: 1-509. Paris. Girt, T. 1863. Synopsis of the family of Lepturoids, and description of a remarkable new generic type. Proc. Acad. nat. Sci. Philadelphia, 1863 : 224-229. Goopg, G. W. & Bran, T. A. 1895. Oceanic Ichthyology. Spec. Bull. U.S. nat. Mus, 2: I-553, appendix, figs., pls. Goopricn, E. S. 1909. Vertebrata Craniata: Cyclostomes and Fishes. In A Treatise of Zoology, ed. Ray Lankester, 9: xvi + 518, figs. Grecory, W. K. 1933. Fishskulls: A study of the evolution of natural mechanisms. Tvans. Amer. Phil Soc. 23 : vii + 75-481, figs. Grey, Mrs. M. 1953. Fishes of the Family Gempylidae, with records of Nesiaychus and Epinnula from the Western Atlantic and descriptions of two new subspecies of Epinnula orientalis. Copeia, 1953 : 135-141. GuntuErR, A. 1860. Catalogue of the Acanthopterygian Fishes in the collection of the British Museum, 2: 1-548. London. Jounson, J. Y. 1865. Description of a new genus of Trichiuroid Fishes obtained at Madeira with remarks on the Genus Dicrotus Giinther and on some allied genera of Trichiuridae. Proc. zool. Soc. Lond. 1865 : 434-437. Jorpan, D. S. 1923. A classification of Fishes, including Families and Genera as far as known. Stanford Univ. Publ. Biol. Ser. 3 No. 2 : 1-243, i-x. Martsupara, K. & Iwatr, T. 1952. Studies on some Japanese fishes of the Family Gempylidae. Pacif. Sci. 6 : 193-212, figs. Route, L. 1927. Considérations sur deux espéces abyssales entrées au Musée Océanographique et sur la valeur tératologique possible de l'une d’elles. Bull. inst. Océan. Monaco, 497 : I-12, figs. Recan, C.T. 1909. On the anatomy and classification of the Scombroid Fishes. Ann. Mag. nat. Hist. (8) 3 : 66-75, figs. Srarks, E. C. 1911. Osteology of certain Scombroid Fishes. The Osteological characters of the Scombroid fishes of the Families Gempylidae, Lepidopidae and Trichiuridae. Stan- ford Univ. Publ. No. 5 : 17-26, pls. STEINDACHNER, F. 1867. Ichthyologischer Bericht iiber eine nach Spanien und Portugal unternommene Reise. IV: Ubersicht der Meeresfische an den Kiisten Spaniens und Portugals. Sitzb. K. Acad. Wien. 56 : 1-106, pls. Swainson, W. 1839. The Natural History of Fishes, Amphibians and Reptiles, 2 : 1-452, figs. (in Lardner’s Cabinet Cyclopaedia, London). Tucker, D. W. 1953. The Fishes of the Genus Benthodesmus (Family Trichiuridae). Proc. zool. Soc. Lond. 123 : 171-197, pls., text-figs. (‘yjIeg ‘umasnyy UeTpeAJsNY 4saA\ ay} JO OJIN ‘AonepH “T “Ac jo Asoyinos Aq ydersojoyq) “Typ ‘wur S11 ‘odAjzojoxY jo pray *(tapuexely) 9vzwy sasans GIL Ny Acl "€ ‘H'N) “Wd ‘1g | ee “RELATION SEUPS. AND _ EVOLUTION CELE SUPER SPECIES (AVES, MUSCICAPIDAE) _ BULLETIN OF | Vol. 4 No. 4 LONDON: FOgGay VARIATION, RELATIONSHIPS AND EVOLUTION IN THE PACHYCEPHALA PECTORALIS SUPERSPECIES (AVES, MUSCICAPIDAE) BY IAN C. J. GALBRAITH Ph. 131-222 ; 8 Text-figures. BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 4 No. 4 LONDON: 1956 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, 1s issued in five series corresponding to the Departments of the Museum, and an Historical Series. Parts 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. This paper is Vol. 4, No. 4 of the Zoological series. PRINTED BY ORDER OF THE TRUSTEES OF THE BRITISH MUSEUM Issued September, 1956 Price One Pound Four Shillings VARIATION, RELATIONSHIPS AND EVOLUTION IN” THE |/PACHYCEPHALA PECTORALIS SUPERSPECIES (AVES, MUSCICAPIDAE) By IAN C. J. GALBRAITH SYNOPSIS The Australasian species Pachycephala pectoralis is remarkable for the great number and variety of geographical representatives which, since they intergrade, must be included in it. Although cited in recent evolutionary literature (Meise 1936, Dobzhansky 1937, Mayr 1942, Ripley 1945, Cain 1954a), the species has not previously been revised as a whole. It presents a wealth of geographically-variable plumage characters, whose relative systematic importance can be assessed from their co-variation. The complicated character-geography of the P. pectoralis superspecies is here interpreted in terms of colonizations by two major stocks, followed by divergence in isolation, great expansions of range, and extensive secondary intergradation. Whether two forms, on meeting, will interbreed or coexist as distinct species, seems to depend less on their degree of relationship than on internal and external ecological factors. CONTENTS Page INTRODUCTION . 5 : 6 2 4 5 . 2 rels5 Scope and presentation c 3 5 : c . 0 5 = 130 Material B 5 3 : - : : 5 : ol WEY/ PLuMAGE PATTERNS . ; 4 a 0 ; 5 : 6 c iey! Pigments. 3 0 . = 5 2 é . . 6 EY Patterns : 7 ¢ : : : 5 teats} Retarded and juv ile plumages “ c a . A 5 so Visual significance : z c ¢ < c c c 39 VARIATION F > 3 6 : © 5 6 cj : - 140 Size variation : zi 5 : - 5 ei = 40 Male character- aeeopciphy : c c : é c 0 Jae Female character-geography . ; 3 4 4 5 c LAS NaturAL Groups 3 : @ : x = A'S, Co-variation and character-complexes : c : : - 148 Intergradation F 5 2 F : a : 5 2 etek) Subspecies-groups . c 5 : - 4 2 ¢ 5 - 149 P. schlegelit . : o B 6 : 4 5 : - 150 Lesser Sundan group A . : : : 3 ° - 150 Moluccan group B 4 4 : 0 : . : 3 5, aise Solomons group C . P ° - n : 0 , : a EGE Fijian group D 1 ° : A A : : ° = 153 P. flavifrons . é : 6 5 r “ ¢ = 54 P. soror a P 0 A é » 54 Northern Australian group i 5 P : é c 4 Southern Australian group F . 4 c : 4 . c 50) Southern Melanesian group G . - : 5 7 : 5 5 7 Widespread groupH . A e 4 e 5 é ° - 158 ZOOL. 4, 4. 10 THE PACHYCEPHALA PECTORALIS SUPERSPECIES CONTENTS—cont. Page INTERGRADATION AND BRIDGELESS GAPS . . . . A + 160 Gene-exchange between ait N . . : 5 - 160 Marks of ey, a . : : C : : 2 Os Sympatry . 5 : : . 3 3 : 5 - 166 Group AFFINITIES A 5 ° é d b : c =) 268 The schlegelit Basemiblage : . . . . é LOS: The soroy assemblage. . A 5 : c . 0 + 170 ARRANGEMENT . é 5 5 A é b 3 6 Bly The superspecies . ste tas : 5 : : 4 : = D7 Species : : ° . . : : . . rez Subspecies-groups . : 2 5 4 3 : ¢ 2 LTS Subspecies . v : 2 : 0 c 4 : = EVvoLurTIon . 2 3 ° * F 2 7, Geographical speciation in New Guinea é 3 5 0 . a e/7/ Origin of the dichotomy c 5 0 0 2 5 EGS Development of isolating mechanisms c : : 2 : 7 £79: A suggested course of events . : c 6 = ( 3 = XSt Rate of divergence R ¢ g 0 5 0 5 : -) CESS Unexpected uniformity . 4 = a - 9 b a Los Unexpected diversity . 6 . A , : 3 - 184 ADAPTATION : 6 0 2 3 : : ¢ c - 186 Character and climate E 7 6 “ 0 i A : - 186 Character and habitat . 3 5 a : C : - 188 Variation in habitat 6 5 F 6 7 " 3 5 litsie) Variation in bill size . : : d 5 4 5 a) EOE CONCLUSIONS 6 : 4 a a 0 3 : . 7 On SUMMARY . ¢ G 6 : . cl : : ; +) So” APPENDIX CHECKLIST . : 5 : 5 4 : . 1095 Subspecies to be recornized “ : c 5 a : ° + 204 Notes 5 : - 6 . A 0 : 2205 P. schlegelit viridipectus . Ee 5 . : o 3 . + 205 P. pectoralisjubilarui 2 ° : ; 3 5 . + 206 P. pectoralis atromaculata c : : 5 0 : : - 206 P. pectoralis gilolonis . c : 0 : 5 c : - 206 P. pectoralis ambigua. ‘ A 5 : : 2 s + 206 Female of bynoei . 9 é 5 o - 206 Intergradation of spinicauda with dahli 5 : : c . - 206 The subspecies on Teste Island : 2 ‘ a : 5 + 206 P. pectovalis fergussonis . 3 5 C : a < : ZO, P. salomonis . : 5 5 5 oe White-throated Pachyce pata in the Solomons “ 6 6 : - 208 P. pectovalis brunneipectus, banksiana and efatensis . c 5 - 208 P. macrorvhyncha eee: 5 6 a a 0 2 bu - 208 P.collaris . S 2 ° 9 0 e b dhe P. pectovalis misimae_ . : 0 5 5 : 7 : - 209 Saxicola merula . A : ‘ é 5 3 5 * «200 ACKNOWLEDGEMENTS . F é 0 * 2 : 5 7 - 209 MEASUREMENTS . : 2 2 a o z 2 c - a) ZL Weight : : : 5 - : 5 ; 5 Gein Wing and tail lenpeas e 0 a é 0 Sein Tarsus and culmen lengths, and bill ‘depth 2 . ° : eae REFERENCES : S cd F : : 5 . a een THE PACHYCEPHALA PECTORALIS SUPERSPECIES 135 FIGURES Page 1. Standard plumage patterns 5 c : . - : c c ws¥s} 2. Extreme forms of P. pectoralis . j 6 5 6 - 0 - I61 3. Sympatric species in New Guinea 2 c - c 6 5 a BS 4. Diagram of relationships . c 0 : + 169 5. P. flavifrons, P. p. melanops and P. i. ] caledonica 6 o 0 173 6. Sketch-map of distribution : 0 6 a end-fold 7. Diagram of colonizations . : : : 5 2 5 . end-fold 8. Diagram of secondary intergradation . - ; 0 5 . end-fold TABLES Page I. Composition of plumage colours. ¢ C : : : Ss 7 II. Colour-phases in P. flavifrons . : . 5 6 0 . 154 III. Characters uniting the schlegeliz sesoninene : o 39/0) IV. Characters uniting the soror assemblage (except subspecies: group H) . 171 INTRODUCTION Pachycephala pectoralis, the Golden Whistler, is probably unique in the richness of its geographical variation. More than seventy subspecies can be recognized, extending from Java and the Moluccas to Tasmania and Tonga. Many of these are so unlike that they would certainly be considered as distinct species, were it not for the more or less complete intergradation between them. There has been no comprehensive checklist of the species since that proposed by Mathews (1930), in which forms now generally recognized as subspecies of P. pectoralis are separated into eleven species. But as early as 1908 Rothschild and Hartert had suggested that the distinctive races of the Solomons are conspecific with more characteristic P. pectoralis, and this was confirmed twenty years later (Hartert, 1929) by the discovery of hybrid populations. Rensch (1931) included the Sumban race and its relatives, but not those of the northern Moluccas. Mayr (1932a, 6) placed even the aberrant forms of northern Fiji in P. pectoralis, because they are connected with that species by intermediate forms. The conspecificity of diverse forms has been accepted in subsequent lists (van Bemmel, 1948 ; Mayr, 194Ia, 1945, 1954a, 1955), which together cover almost the whole range of the species. All these forms are accepted here as belonging to P. pectoralis. P. soror in the hill forest of New Guinea is very like nearby races of P. pectoralis. Over most of its range it replaces this species; but a race of P. pectoralis lives so close to populations of P. sovor (Rand, 1940) that the barriers between them are probably intrinsic (p. 166). Thus two very similar forms seem to be genetically sym- patric (Cain, 1953), and must be considered as distinct species ; while others which differ much more, and seem to be actually less closely related, intergrade and must be considered conspecific. This situation is not unknown in other animals (e.g. Acanthiza, Mayr, 1942, 174), but the striking example in Pachycephala has been pointed out only by Cain (19542) as a result of the present review. 136 THE PACHYCEPHALA PECTORALIS SUPERSPECIES Scope and presentation Four species are considered, which form a single superspecies with a triplet (see p. 172) in New Guinea: P. schlegelit Schlegel, New Guinea mountain forest from 4,000 or 5,000 to 12,000 ft. P. sorory Sclater, New Guinea hill forest from 2,200 to 5,200 ft. P. pectoralis (Latham), Lesser Sunda Isles and Moluccas to Tasmania and Tonga; but absent from New Guinea except for the south-east coast, and disturbed habitats in the Snow Mountains between 5,200 and 8,000 ft. P. flavifrons (Peale), Samoa. No linear arrangement can be satisfactory, since P. pectoralis connects the other three species. Text-figure 6 (end-fold) gives the ranges of the species and subspecies, and of the subspecies-groups of P. pectoralis. The latter seem to be natural groups, though because of gene-interchange their boundaries are not sharp and have to be shown rather arbitrarily (see Text-fig. 8): Lesser Sundan subspecies-group A (subspecies I-5). Moluccan subspecies-group B (subspecies 6-8). Solomons subspecies-group C (subspecies 9-17). Fijian subspecies-group D (subspecies 18-21). Northern Australian subspecies-group E (subspecies 22-27). Southern Australian subspecies-group F (subspecies 28-33). Southern Melanesian subspecies-group G (subspecies 34-38). Widespread subspecies-group H (subspecies 39-57). P. schlegelii and P. soror have three subspecies each, while P. flavifrons is mono- typic. Many forms recognized as distinct subspecies by recent authors have been combined in this presentation (see p. 175), although a number of these are distinct enough to be separated according to current usage (list on p. 205). The use of subspecies names is not helpful to the reader unless he is already familiar with the group under discussion. Nor is it usually necessary, since most subspecies are easily characterized by their geographical ranges (cf. Wilson & Browne, 1953). In this paper the range citation, given in a condensed and approximate form, is followed by a cipher for direct reference to the map (Text-fig. 6). This cipher consists of the number of the subspecies within its species, preceded by the subspecies-group letter for subspecies of P. pectoralis. Where infrasubspecific variation is discussed, parts of the subspecies range are indicated by lower-case suffixes. Thus “Sumbawa to Alor A3’’ indicates the form of P. pectoralis on Sumbawa, Flores, Lomblen, Pantar and Alor (which from “‘Lomblen to Alor A3b”’ is slightly larger and larger-billed). The exact range of any form can be found from the check- list (p. 195), which is lettered and numbered to correspond. Subspecies names are useful in referring to forms whose geographical ranges are diffuse or difficult to define: “ dahli E25’ is used for the subspecies of P. pectoralis which ranges from south-eastern New Guinea and Fergusson Island to many small islands in the Bismarck Archipelago. The range of dahli is shown inset on the map. THE PACHYCEPHALA PECTORALIS SUPERSPECIES 137 Material I have seen at least one adult male and one adult female of every subspecies recognized by recent authors, except for the following: no specimens of H46 and H50; no adult male of D21; no adult female of C15, G37a & b, H49, H52 and H56 (juveniles seen) and E25c (female unknown). I have examined the following types in the British Museum (Natural History): fulvotincta Wallace A3a, mentalis Wallace Bo, neglecta Layard (= D18b), aurantiiventris Seebohm Diga, torquata Layard D2o0a, fuliginosa Vigors & Horsfield F28b, fusca Vigors & Horsfield (= F30b), variegata Gray (= G34), cucullata Gray G36, chlorurus Gray G37a, intacta Sharpe G37d, fuscoflava Sclater H42, xanthocnemis Gray (= H43b), clio Wallace H45, collaris Ramsay H47a, vitiensis Gray H55, klossi Ogilvie-Grant and bartoni Ogilvie-Grant (P. soror 2 & 3). Unfortunately, many of the available series were very short, and the measurement tables compiled (p. 212) are inadequate for proper statistical treatment. Individual variation and fine geographical variation are therefore not considered in this paper. Where the available material was inadequate full use has been made of published descriptions and measurements, especially those of Mayr (1932a, b, and 1954a). I have studied P. pectoralis in the field on Guadalcanal C11 and San Cristobal C17a, and seen and heard it near Sydney F30b, on Lord Howe and Norfolk Islands F32 & 33, and on Efate and Santo G37b & d. I have also studied P. implicata (which is rather closely related to the pectoralis superspecies) in the mountains of Guadalcanal (Cain & Galbraith, 1956). PLUMAGE PATTERNS Pigments All the colours in the various plumages of the superspecies are produced by combinations of yellow, black and brownish pigments (Table I). The yellow pigment is soluble in boiling alcohol or pyridine, and turns a transient blue-green with con- centrated sulphuric acid. It is therefore a carotenoid (Cain, 1950, 104). Carotenoid TABLE I.—Composition of Plumage Colours. Carotenoid. Melanins. None. V. pale. Pale. Deep. None . A 5 : > White Pale yellow Lemon-yellow Golden- to orange-yellow Eumelanin : Barbs clear, barbules satu- Grey Olive-grey Olive-green Golden-olive tated Granules in barbs, barbules Dull black ae Olive-black — saturated Saturated . 0 : - Black —_ — — Phaeomelanins : Pale . 5 6 : - Sandy to Cream Cinnamon Citrine dull pink to vinous Deep . 5 . - Brown to Russet Tawny-orange — Tufous 138 THE PACHYCEPHALA PECTORALIS SUPERSPECIES tends to be more concentrated in the barbs than in the barbules of the feather. The other pigments, if not saturated in both elements of the vane, are more concentrated in the barbules than in the barbs. They are granular. Under the microscope black granules still appear intense black, while brown ones vary considerably in hue and intensity. The black and brown pigments are presumably eumelanins and phaeome- lanins. Parts of the vane devoid of melanins are filled with minute bubbles, which by multiple internal reflection add to the brilliance of the whites and yellows. Most contour feathers are grey basally, with the black granules forming bands across the barbules. Patterns It is convenient to describe the variation of the male pattern throughout the superspecies in terms of departure from a standard. The descriptions will be briefest, and the peculiarites of the various forms most clearly apparent, if the pattern chosen as standard combines all the variants which are more common than their alternatives. This condition is fulfilled by the pattern of males of P. pectoralis in the Bismarcks H48-51, which is shown diagrammatically in Text-fig. 1 (top), and described below. Fic. 1.—Diagram of the standard plumage patterns for the superspecies. Male (above, facing left) H48-51 ; female (below, facing right) H52. Not to scale. Colour key : diagonal hatching, grey (without stipple) or olive (with stipple) ; cross-hatching, brown and rufous hues; stipple, yellow ; black, black; white, white. The same conventions are used in all the diagrams of plumage pattern (Text-figs. 1-3 and 5). ADULT MALE. White chinand throat separated by black gorget, between auriculars, from golden-yellow underparts; fore-edge of gorget formed by black tips of white feathers, hind-edge by black feathers overlying yellow; black cap and auriculars separated by yellow collar across hind-neck from golden-olive mantle ; wings dull black with olive outer edges to all feathers, fading to grey towards tips of outer primaries ; inner edges of quills whitish ; tail and upper tail-coverts black with olive tips, and olivaceous traces at bases of outer quills. It is harder to select a standard for the female plumage, since the patterns are much vaguer than those of the males, and the variation is more quantitative than qualitative. The following description and the diagram (Text-fig. 1, bottom) best fit some females of the widespread group H, and especially those from Ndeni H52. ADULTFEMALE. Throat pale buff with faint dark fringes at sides ; gorget brownish, THE PACHYCEPHALA PECTORALIS SUPERSPECIES 139 vaguely defined, grading into brownish wash on breast and flanks ; mid-belly and under tail-coverts free from melanins; under tail-coverts pale yellow, breast and belly washed with yellow ; upper parts citrine ; collar, rump and tail brighter ; cap greyer than mantle, auriculars cinnamon ; wings blackish-brown with citrine outer edges to all feathers, fading to greyish towards tips of outer primaries ; inner edges of quills whitish. Retarded and juvenile plumages Males are sometimes found in plumage like that of the adult females (‘‘ II Phase ”’ of Mayr, 19322, b), though commonly with more intense carotenoid. They agree with adults in tail shape and in having black bills, and their testes are often enlarged (Baker, Marshall & Harrison, 1940). They are evidently adult males in a “ retarded ” plumage (Mayr, 1933): in Victoria and Tasmania at least they are known to breed in this condition (Chandler, 1912 ; Howe, 1927 ; Lawrence, 1952). Juveniles of both sexes have brown or straw-coloured bills, and softer wings and more pointed tail-feathers than adults. In pattern they resemble adult females, except that they have the edges of the wing feathers rufous, the cap olivaceous, the mantle often browner and the carotenoid pigment more dilute (I Phase). Nestlings are commonly rufous all over ; but in the lesser Sunda Isles (nestling of A3b seen) are greyish with darker shaft-streaks beneath. Only Mayr (1932a, b) has systemati- cally described immature plumages, and I have not examined sufficient immature skins from parts of the range not treated by him to attempt a survey of their varia- tion. At first sight the female and juvenile patterns seem very different from that of the adult male. But the distribution of pigments is much the same—except that carotenoids are greatly diluted, there are no solid blacks, and eumelanins are more or less replaced by phaeomelanins. As a result, these patterns are less sharply defined and stereotyped than that of the male, and show a greater range of colours. Visual significance In many birds the upperparts are cryptic, while the underparts bear a conspicuous pattern which plays a part in intraspecific display. The standard male pattern of P. pectoralis is a good example of this. Both the ventral and the dorsal pattern embody transverse bands which contrast strongly with the ground-colour (see Cott, 1940). The bright underparts contrast with the forest background, so that the black gorget (exploiting maximal tone-contrast against the white, and an especially effective colour-contrast against the yellow) is an emphatic feature. The olive upperparts, on the other hand, conform with the background, and the yellow collar is disruptive in effect. Females and juveniles tend to be inconspicuously coloured, with neither emphatic nor markedly disruptive features. This common species has been largely neglected by Australian ornithologists (except Lawrence, 1952), and nothing seems to be known of the part played in intraspecific display by the conspicuous male ventral pattern. Although we saw neither courtship nor aggressive displays in the Solomons, the general behaviour of 140 THE PACHYCEPHALA PECTORALIS SUPERSPECIES these birds (Cain and Galbraith, 1956) is much what might be expected from a study of their colour-patterns. The males show themselves freely to a ground observer as they move about below the canopy, whistling loudly. But although they will sing from bare branches overhung by foliage, they avoid all perches exposed to the sky. Females of P. pectoralis on Guadalcanal C11, like those of many other subspecies, are dull in colour. They are very silent and skulking, seldom seen as they move about in dense thickets near the ground. Those on San Cristobal Cr7 are as bright yellow beneath as the males, but lack the black gorget. They are often seen in company with the males in the upper substage, usually silent, but occasionally heard to give alarm calls and quiet whistles. A related species in the mountains of Guadalcanal (P. implicata) has the females actually more conspicuous than the males. They are not more retiring than the males, and are more vocal than the females of P. pectoralis. The males of P. implicata are inconspicuously coloured in olive and blackish, and are much more silent and retiring than those of P. pectoralis. On the other hand, a pair of this species was seen feeding exposed in the tops of small trees: perhaps the direction of predation is less important to these more uniformly-coloured birds. VARIATION Geographical variation in the superspecies is conspicuous and complex. Marked sexual dimorphism is normal, but in three widely-separated races of P. pectoralis the males are permanently hen-feathered, while P. flavifrons has cock-feathered females. The sporadic occurrence, in sexually dimorphic forms, of breeding males in female plumage shows that hen-feathering is not very important here, either as an indicator of genetical change or as a potential barrier to interbreeding. The bold pattern of the male is subject to striking variation. For example, in some forms the throat is yellow instead of white, and some of these lack the black gorget. These differences are not unifactorial, since for almost every pair of contrasted characters intermediate states are to be found. Single characters cannot be used to delimit natural groups, since not one variant is confined to a single subspecies-group of P. pectoralis (as distinguished by the whole constellation of characters of both sexes) and shown by every race belonging to that group. Most are both polyphyletic in origin and labile within groups. The marked geographical variation of the female pattern does not present the same appearance of discontinuity, and there is less temptation to erect a classification based on single characters of the females. Size variation The largest subspecies of P. pectoralis have wings over 30% longer than the smallest. If their proportions are the same, they should therefore be more than twice as heavy. Unfortunately, weight measurements are available for very few forms in the super- species, so that wing lengths are for the present the best indication of body size. Certain subspecies which are evidently very closely related differ markedly in size, which is therefore not generally of much importance in assessing relationships ; but some of the discontinuities between subspecies-groups are marked by sharp changes THE PACHYCEPHALA PECTORALIS SUPERSPECIES 141 in size as well as in other characters. A coarse grading into size-classes adequately expresses the major variation, which more detailed treatment would tend to obscure. In any case, too few specimens of many forms were available for quantitative treat- ment to be satisfactory. Snow (1954) has pointed out that quantitative differences (in size, proportions, intensity of pigmentation, etc.) are often adaptive, and cannot be used as clues to relationships and past history until their present environmental relations have been worked out. Conversely, where the past history has been as com- plex as in the fectoralis superspecies, such differences cannot be shown to be adaptive until relationships have been established (by studying the co-variation of more stable characters). Two forms on neighbouring islands might differ in size, not because the climates were different but because they were the same, if the forms belonged to different subspecies-groups whose size/temperature relationships (Snow, 1954, 22) had been evolved under different climates. The various forms are graded into five classes on the average wing lengths of adult males. But the arbitrarily-selected limits of these classes have in a few cases been relaxed, to avoid separating closely-related forms whose small difference in size happens to transgress them. The southernmost Australian forms are classed as “large”, not as “ very large’’, because the appearance of the skins (prepared by many different collectors) strongly suggests that they are considerably smaller birds, though with relatively long wings (perhaps in relation to their nomadic habit) than the ‘‘ very large’ ones of the Solomons and elsewhere. This can only be decided when weight measurements are available. The middle class, containing the most forms, is of wing lengths from 90 to 95 mm.; those from 80 to 85 mm. are graded as “very small’, from 85 to go as “‘small’’, from 95 to 100 as “large ” and over 100 as “very large’. The following forms fall outside the middle class : VERY SMALL BIRDS. Sumba, Java to Alor, and Flores Sea and Salayer A1-5 ; midwestern Australia E22 a—c (size increasing north-eastwards). SMALL BIRDS. Rennell and San Cristobal C16—-17 ; northern Australia to southern New Guinea E2zd-24 ; New Caledonia, New Hebrides and Vanikoro G34 & 36-38 ; Timor to Damar H39-41; Sula and Peleng Isles H45~-46; large islands in the Bismarcks H48; Kandavu H54; P. soror; P. schlegeli; P. flavifrons. LARGE BIRDS. Ternate, Tidore and Obi B7-8; Vella Lavella to Gatukai C13 & 15; Tasmania and southern Australia F28-30, size decreasing northwards ; Tabar H50; Lau archipelago H56. VERY LARGE BIRDS. Bougainville to Malaita, and Rendova and Tetipari Cg-12 & 14; Loyalty Isles G35; Tenimber Isles H42; Lihir H49; Tonga H57. Differences in bill size are even more marked, with the largest bills some 60% longer than the smallest, and the bill may be stubby, stout or slender. However, closely related forms differ in bill size even more than in wing length, and the accuracy of this measurement is not sufficient for the forms to be graded with any degree of certainty from the scanty data available. Tail and tarsus lengths are only comparable when expressed as relative lengths, which vary between individuals much more than the absolute measurements. Therefore none of these measurements is dealt with comprehensively, though there is some discussion of climatic correlation in certain parts of the range (p. 186). 142 THE PACHYCEPHALA PECTORALIS SUPERSPECIES Male character-geography Some of the most conspicuous variations appear to be qualitative and discon- tinuous, but hybrid forms often show intermediate states. Probably most of the variations are in fact potentially continuous—but it is convenient to deal with the geographical distribution of the more important variations in terms of discontinuous “characters’’, at the same time mentioning the occurrence of intermediate conditions. (1) HEN-FEATHERED. Rennell C16, Norfolk Island F33 (sexes almost indis- tinguishable) ; Salayar A5 (male in a female type of plumage, but more advanced in character than its own female). The hen-feathered races were considered as separate species until Mayr (19322, 5) pointed out the relative unimportance of the character. Since individual males of dimorphic subspecies can breed in female plumage, hen-feathering involves neither a radical change in the genome nor complete breakdown of courtship and territorial displays. Variations of the male pattern such as “no gorget’”’, “no collar’’, “ cap pale’’ and “‘ tail pale’ may be considered as partial hen-feathering, and the San Cristobal form C17 especially shows several such indications. The three fully hen-feathered forms are not considered further in the male character- geography. (2) CHin BLACK. Sumba Art; Morotai to Batjan B6-7; Solomons except San Cristobal Cg-15 ; P. schlegeli1, P. flavifrons. (3) THROAT AND CHIN BLACK. Tonga H57. In P. flavifrons there is much black at the base of the throat feathers, which shows as irregular barring. Another approach to the black-throated condition is shown in the reduction of the throat patch on Sumba Ar (where black malar feathers intervene between it and the auriculars), from Java to Alor A2-3, and in P. soror and P. schlegelit. (4) THROAT YELLOW. Solomons Co-17 (variable in hybrids on small islands off Shortland E26) ; northern and central Fiji Dr8-21 ; Tenimber Isles H42; P. flavi- frons (individually variable). Juveniles on Vanikoro G38 have yellow throats, and there is a tendency for the throats of juveniles in the New Hebrides G37 and on Utupua H53 to be yellowish (Mayr, 1932b). A few yellow feathers occasionally appear in the white throats of adult males. I have seen these in males of P. pectoralis from Teste Island E25b, Lord Howe Island F32, New Caledonia G34, Loyalty Islands G35, Aneitum G36 and Kandavu H54, and in P. soroy from the Snow Mountains 2a. But they seem to occur most frequently in P. schlegelii—in eight of thirty-one adult males examined. (5) BASES OF THROAT-FEATHERS WHITE. Obi B8; Solomons Cg-17; northern and central Fiji Dr8-21 ; midwestern Australia to small islands in the Bismarcks, and hybrids off Shortland E22-26 ; Vanikoro G38; Tenimber Isles H42 ; Sula and Peleng Isles H45-46; Louisiades H47 ; Ndeni, Santa Cruzislets and Utupua H52-53. (6) No GorcetT. Malaita C12; Viti Levu and Vanua Levu D18a and 1ga; P. flavifrons. The gorget is more or less broadly interrupted in individual males on the islands between Koro and Taviuni on the one hand, and Viti Levu and Vanua Levu on the other—D18b, rgb & c. THE PACHYCEPHALA PECTORALIS SUPERSPECIES 143 (7) FORE-EDGE OF GORGET DEFINED BY PALE FEATHERS. Sumba and Java to Alor A1-3; northern Moluccas B6-8; Solomons except San Cristobal (and the gorgetless forms on Malaita and Rennell) Cg-11 & 13-15 ; Ceram H43; Ndeni and Santa Cruz islets H52 ; P. schlegelit. In these forms the black of the gorget deeply undercuts the white or yellow of the throat, whereas in the standard pattern the edge of the gorget is defined on the under- lying layers of feathers at about the same level. On San Cristobal C17, in the hybrids on the outliers of Shortland E26, and on Buru H44, the gorget undercuts the throat- patch, but there are also some black tips. (8) GORGET CUT OFF FROM AURICULARS. Northern Moluccas B6-8 ; New Caledonia G34; Ceram H43; Ndeni and Santa Cruz islets H52. In all these forms the throat-feathers are unusually long, extending further back than the auriculars. They are best developed from Morotai to Batjan B6-7 (where they cover most of the gorget), and least so on New Caledonia G34 (where only the lateral feathers are appreciably lengthened). In B6-7 and G34 the gorget is itself reduced, and does not join the auriculars even beneath the white feathers. On Vella Lavella and Ganonga C13 the yellow feathers extend behind the auriculars, but the gorget is too broad to be cut off. (9) GORGET VERY BROAD. Vella Lavella and Ganonga C13; P. schlegelii. (10) GORGET VERY NARROW. New Caledonia G34; Kandavu H54; P. soror. (11) BREASTRUFOUS. Sumba Ar (intense, extending to flanks and belly), Java Aza (less intense, especially on belly), Bali Azb (still less intense on flanks and belly), Sumbawa to Alor A3 (confined to band behind gorget), islands in Flores Sea A4 (very faint, barely detectable on belly and flanks) ; Santa Anna Cr7b and individuals on San Cristobal C17a (patch behind gorget); New Caledonia G34 (patch behind gorget) ; P. schlegelit (extending to flanks and belly; very intense from Snow Mountains to south-eastern New Guinea 3, somewhat less so in Vogelkop and Cyclops Mountains 1 & 2). There is a rufous wash centred on the vent on Taviuni, Koro and Vatu vara D2o-z21, Ceram H43, Ngau H55, and in individuals on Aneitum G36. Where very pale rufous phaeomelanin combines with pale yellow carotenoid (as in the Flores Sea A4), the tawny-orange produced is rather like the orange-yellow of very intense carotenoid (as on Vanua Levu Drg). The difference is easily seen through the microscope, since the phaeomelanin appears granular. (I2) CAROTENOID VERY DEEP (ORANGE-YELLOW). Vella Lavella and Ganonga C13; Vanua Levu to Koro and Vatu vara Drg-21 ; Louisiades H47; Lihir H49; Manus H5r1. (13) CAROTENOID PALE (LEMON-YELLOW AND GREENISH-OLIVE). Mid- and north- western Australia E22, deepening northwards E22a to 23; Tasmania and southern Australia F28~30, deepening northwards F29 to 31 ; New Caledonia, New Hebrides and Vanikoro G34 & 36-38; Babar H4o; Utupua H53; P. soror. (14) ForREHEAD YELLow. Viti Levu, Vanua Levu and Vatu vara Dr8a, Iga & b & 21 (sometimes a few yellow feathers on Ovalau, Rambi and Kio D18b & 19Qc) P. flavifrons (sometimes white, in white-throated individuals). (15) CaP PALE. Santa Anna C17b (olive, with blackish lores and auriculars) and > 144 THE PACHYCEPHALA PECTORALIS SUPERSPECIES San Cristobal Crza (variable, from olive to black) ; New Caledonia G34 (grey) ; Aneitum G36 (dull black with more or less olive scalloping). (16) No cottar. Vella Lavella and Ganonga C13, Ndeni and Santa Cruz islets H52, P. flavifrons (melanic forms with no trace of a collar) ; Fiji, except for Vatu vara and Kandavu, D18—20, H55-56 (black of cap extends far down on hind-neck, followed by a yellowish trace) ; Malaita C12, New Caledonia G34, P. soror (yellowish trace) ; San Cristobal C17 (yellowish trace on hind-neck, clear yellow patches at sides of neck). The collar is very vague and narrow on the islands in the Flores Sea A4, from Kulambangra to Gatukai C15 and on the Tenimber Islands H42. It is narrow and washed with olive on the nape on Sumba and from Java to Alor A1—3, on Rendova and Tetipari C14, in mid- and north-western Australia E22, in Tasmania, southern and eastern Australia and on Lord Howe F28-32, from the Loyalty Isles to Vanikoro G35-38, on Timor H39a, the Louisiades H47, Utupua H53 and Kandavu H54. (17) MANTLE BLACK. Vella Lavella and Ganonga C13, Ndeni and Santa Cruz islets H52, P. flavifrons (wholly black) ; islands in Flores Sea A4, Rendova and Tetipari C14, Taviuni and Koro D20, Ngau and Lau archipelago H55~-56 (feathers black-centred, individually variable from olive with concealed black spots to black with olive scalloping). (18) WinG BLack. Islands in Flores Sea Aq; Vella Lavella, Ganonga, Rendova and Tetipari C13-14; P. flavifrons; P. schlegelit. In these forms the pale edges to the wing-feathers are absent or exceedingly narrow. The pale edges are narrow, and the primary-coverts entirely black, on Taviuni, Koro and Vatu vara D20-21, in dahli E25, on Ceram and Buru H43-44, Ndeni and Santa Cruz islets H52 and Ngau and the Lau archipelago H55-56. (19) OLIVE VERY DARK. Kulambangra to Gatukai C15 ; Fiji Di8—21 & H54-56 ; Ceram and Buru H43-44; P. soror; P. schlegelit. Normally the bases of the barbules, as well as the barbs, are yellow, giving a more or less pronounced herring-bone pattern under the microscope. In the above forms the barbules are black down to their junctions with the barbs. (20) OLIVE TINGED WITH BROWNISH. San Cristobal C17; Koro and Taviuni D20; Ceram H43; Louisiades H47; individuals on Aneitum G36. The phaeomelanic wash is deepest on tail, upper tail-coverts, rump and secondaries. (21) WING QUILLS GREY-EDGED. Midwestern Australia to small islands in the Bismarcks E22-25 ; Tasmania F29; Damar H41; P. soror in Snow Mountains 2a. The wings are unusually grey in southern Australia F28 & 30, becoming more olive northwards ; and in eastern Fiji Dig & 21 & H56, becoming more olive westwards (towards Viti Levu and Kandavu D18 & H54). They are also unusually grey in all populations of P. soror. They are variable in colour in the hybrids on outliers of Shortland E26. (22) UPPER WING-COVERTS YELLOW-EDGED. Viti Levu and Vanua Levu D18—109 ; midwestern Australia to small islands in the Bismarcks E22—25 ; Tasmania, southern and eastern Australia and Lord Howe F28-32; Loyalty Islands G35. The upper wing-coverts are edged with yellower olive than the mantle in many forms ; those listed above show this most conspicuously. THE PACHYCEPHALA PECTORALIS SUPERSPECIES 145 (23) UPPER TAIL-COVERTS ALL-OLIVE. Kulambangra to Gatukai C15; San Cristobal C17; northern Fiji, except Koro and Vatu vara, D18-20a; mid- and north-western Australia E22; Tasmania, southern and eastern Australia and Lord Howe F28-32 ; New Caledonia to Banks Islands G34-37 ; Timor H39; Tenimber Isles H42 ; Louisiades H47 ; Kandavu H54; Tonga H57; P. soror. (24) Tait PALE. Tasmania F29, New Caledonia, Loyalty Islands and Aneitum G34-36, Timor H39, western Louisiades H47a (no solid black) ; San Cristobal C17 (more or less black) ; south-western and South Australia F28, New Hebrides from Erromango northwards G37, Misima H47b, Kandavu H54, P. soror except the Snow Mountains 1 & 3 (black subterminal patch) ; south-eastern Australia and Lord Howe F30 & 32, Babar H40, Rossel H47c (more than half black) ; mid-western Australia E22a-c, southern Queensland F31a, Damar H41, P. sorory in Snow Moun- tains 2 (wide olive edges basally) ; north-western Australia E22d, northern Queensland F3rb (narrow olive edges basally). The pale part of the tail, usually olive, is grey from south-western Australia to Tasmania and Victoria F28—30a, and sometimes greyish in northern Queensland F3rb. (25) UPPER TAIL-COVERTS ALL-BLACK. Sumba, Java to Alor, and islands in Flores Sea AI-4; Morotai to Obi B6-8; Vella Lavella and Ganonga C13; Ceram, Sula and Peleng Islands H43 & 45-46; Ndeni and Santa Cruz islets H52; P. schlegelit ; P. flavifrons. Even in these forms, the shortest coverts usually have narrow olive tips. Males on Buru H44, and from some other localities, have the tips of all the coverts extremely narrow. (26) TAIL ALL-BLACK. Sumba Ar; Morotai to Batjan B6-7; Vella Lavella and Ganonga C13 ; Ndeni and Santa Cruz islets H52 ; P. schlegelit ; P. flavifrons. There are sometimes narrow and obscure olivaceous tips to the lateral tail feathers, even in these forms. The tips are unusually narrow from Java to Alor A2—3; on Obi B8, in most of the Solomons Cg-12 & 14-15, and from Ceram to Peleng H43-46. Female character-geography (r) COCK-FEATHERED. P. flavifrons (throat and forehead patterns less sharply defined than in the male). Certain single characters of the female (such as a white throat, a sharply defined gorget, bright yellow underparts, a pure grey cap and pure olive mantle, and a partly black tail) tend perhaps towards the male pattern. Often they appear independently, but on San Cristobal C17, and also in northern members of group E and in P. soror, most of the characters mentioned occur together—these forms have an advanced type of female plumage. Individual females of the race on Vella Lavella Cr13a (whose males are all-black above) have black gorgets (Mayr, 1932a, 17). A partly cock-feathered female from Malaita C12 is reported by Mayr (1932a, 21). I have seen several specimens which (though sexed as females) are partly or wholly in the plumage of the adult male, but consider it unwise to rely on the sexing of these, since they carry no indication that the collector had noticed the discrepancy between gonads and plumage. 146 THE PACHYCEPHALA PECTORALIS SUPERSPECIES P. flavifrons is not considered further in the female character-geography. (2) Britt PALE. Solomons Cg-17 (palest from Guadalcanal to Kulambangra and on Rennell Croc—11 & 15-16, darkest on Malaita, Vella Lavella and Ganonga C12 & 13); Rambi and Kio Digc. (3) THROAT WITHOUT MELANIC WASH. Islands in Flores Sea and Salayer A4-5, midwestern Australia E22a-c, New Caledonia to Banks Islands G34-37, Timor and Babar H3 9-40, Louisiades H47, Utupua H53, P. soror (wholly white) ; north- western Australia to small islands in the Bismarcks, hybrids near Shortland and the Snow Mountains E22d-27, Vanikoro G38, P. schlegelii in the Vogelkop and Cyclops Mountains 1 & 2 (white, more or less mottled) ; San Cristobal C17 (pure yellow). (4) THROAT YELLOW. Choiseul to Russel Isles, central Solomons, Rennell and San Cristobal Cro & 13-17; Tenimber Isles H4z. (5) CHEEKS YELLOWER THAN THROAT. Bougainville, Guadalcanal and Malaita Cg, 11 & 12; Viti Levu and Vanua Levu D18a & Iga. (6) THROAT BARRED. Morotai to Obi B6-8 ; Bougainville, Guadalcanal and Malaita Co, 11 & 12; Viti Levu and Vanua Levu D18a & 1ga, and individuals on Ovalau, Rambi and Kio D18b, 1gb-c; north-western Australia to small islands in the Bismarcks, hybrids near Shortland, and the Snow Mountains E22d-27 ; Tasmania, southern and eastern Australia, Lord Howe and Norfolk Islands F28-33 ; Damar H41; Sula and Peleng Islands H45-46; Tonga H57; P. schlegeltt (faintly in the Vogelkop 1). There is no sharp distinction between subterminal barring and terminal fringing of the feathers. There is a tendency towards heavy fringing, not entirely confined to the sides of the throat, from New Caledonia to the Banks Islands G34-37, in the Louisiades and Bismarcks H47-48, and on Utupua H53. (7) THROAT AND BREAST SHAFT-STREAKED. P.p. dahli, hybrids near Shortland, and Snow Mountains E25-27 ; Vanikoro G38 ; Manus H51. (8) UNDERPARTS SHAFT-STREAKED. Solomons except San Cristobal Cg-16 ; Viti Levu and Vanua Levu D18a & Iga, and individuals on Ovalau, Rambi and Kio Di18b & 19b-c. (9) GORGET EUMELANIC. Morotai to Batjan B6-7, P. schlegelii east of the Vogel- kop 2-3 (pure grey); Choiseul and Kulambangra to Florida, Rennell and San Cristobal Cro & 15-17, Loyalty Islands and Vanikoro G35 & 38 (olive) ; Bougain- ville and Malaita Cg & 12, northern Australia E22~23, Tasmania and southern Australia F28-30, New Caledonia, and New Hebrides from Erromango northwards G34, 37a-c & e (considerably greyer than standard). (10) GORGET YELLOW-WASHED. Choiseul to Russel Islands, central Solomons, Rennell and San Cristobal Cro & 13-17 ; Loyalty Islands and Vanikoro G35 & 38 ; Babar and Tenimber Isles H4o & 42 ; Louisiades H47. (II) BREAST AND FLANKS OLIVE. Morotai to Batjan B6—7 ; Choiseul and Kulam- bangra to Florida, and Rennell Cro & 15-16 (pale) ; Snow Mountains E27 (pale, confined to narrow band behind gorget) ; P. soror (pale); P. schlegelit (deep in Snow Mountains and south-eastern New Guinea). Many forms have a combination of carotenoid and phaeomelanins on the under- parts, but this ventral olive is rare. THE PACHYCEPHALA PECTORALIS SUPERSPECIES 147 (12) UNDERPARTS UNIFORM. Koro and Vatu vara D2ob-21, and individuals on Ovalau and Taviuni Dr8b & 20a ; Tenimber Isles H42; Mussau and Lihir H48d—49 ; southern Fiji H54—-56. On Ceram and Buru H43-44 and in the Bismarcks H48a-c, the gorget is little darker than the throat and mid-belly. (13) BELLY WASHED WITH MELANINS. Islands in Flores Sea A4 (pinkish) ; Bougainville, Guadalcanal and Malaita Cg & 11-12 (greyish or rufous buff) ; Rendova and Tetipari C14 (deep rufous) ; Fiji Dr8-21, H54~-56 (greyish on Viti Levu and Vanua Levu D18a & 19a, cinnamon on Koro, Vatu vara and in southern Fiji D2zob-21 & H54-56, individually variable between Koro and the large islands D18b & I9b— 20a); south-western and South Australia F28a-b (pinkish), Tasmania, Victorian mallee and northern Queensland F28c, 29 & 31b (buffy) ; New Caledonia G34 (pale buffy) ; Timor H39 (pinkish) ; Damar, Tenimber Isles, Ceram to Sula Isles, Bismarcks except Tabar H41-45 & 48-49 (brownish, more or less mixed with ellow). (14) BREAST AND BELLY DEEP YELLOW. Morotai to Obi B6-8 (deepest on Obi B8) ; Choiseul and Vella Lavella to Florida, and San Cristobal Cro, 13, 15 & 17; northern Australia, dahli and Snow Mountains E23, 25 & 27; Loyalty Islands and Vanikoro G35 & 38; Babar H4o, Peleng Isles H46, Louisiades H47, Tabar H50, Manus Hs51, Utupua H53, Tonga H57; P. sorvor; P. schlegelit. (15) BREAST AND BELLY SCARCELY YELLOW. Sumbawa to Alor and islands in Flores Sea A3-4; Bougainville, Guadalcanal and Malaita Cg, 11 & 12 (juveniles may be yellower) ; Fiji except Kandavu Dr18-21, H55~-56 ; midwestern Australia E22a-c; Tasmania and southern Australia F28-30; Malekula and Santo G37d ; Timor, Damar and Buru H39, 41 & 44. (16) UNDER TAIL-COVERTS NOT YELLOW. Tasmania and southern Australia F28-30. Here the female plumage is almost devoid of carotenoid, except for a circlet of pale yellow feathers at the vent. (17) Cap ottvE. Solomons Cg-17 (may be obscured by heavy rufous wash). (18) Cap Grey. Salayer A5; Morotai to Batjan B6-7; midwestern Australia to small islands in the Bismarcks E22—25 ; Tasmania, southern and eastern Australia, Lord Howe and Norfolk Island F28-33 ; Vanikoro G38 ; Tabar H50, Lau archipelago and Tonga H56-57 ; P. schlegelii east of the Vogelkop 2-3. (19) AURICULARS GREY. Morotai to Batjan B6-7; Vanikoro G38 ; P. schlegelit east of the Vogelkop 2-3. (20) MANTLE GREY. Tasmania and Victoria F29-30a (wholly grey); south- western and South Australia and New South Wales F28 & 30b (individuals with olive wash on rump or scapulars) ; midwestern Australia E22a—c (rump olive) ; north- western Australia E22d (rump and lower back olive). (21) Tat GREY. Tasmania and southern Australia F28—30. (22) MANTLE PURE OLIVE OR GREY (WITHOUT PHAEOMELANINS). Sumba and Salayer At & 5; Morotaito Batjan B6-7 ; San Cristobal C17 ; midwestern Australia to dahli and Snow Mountains E22-25 & 27; Vanikoro G38; Tabar H50; Tonga H57 ; P. schlegelii in Cyclops Mountains 2. ZOOL. 4, 4. II 148 THE PACHYCEPHALA PECTORALIS SUPERSPECIES (23) UPPERPARTS BROWN. Northern and central Fiji D18-21; Ceram H43, Louisiades H47, Mussau H48d, Lihir H49. The mantle is distinctly brownish-olive on Aneitum and from Mai to Santo G36 & 37c-d, from Timor to the Tenimber Isles H39—42 (tail pure olive on Babar H40), from Buru to the Peleng Isles H44—46, in the rest of the Bismarcks and on Manus H48a—c & 51, on Ndeni, Santa Cruz islets and Utupua H52-53, in southern Fiji H54-56, and in P. soror in the Vogelkop 1. (24) UppERPARTS SOMETIMES RUFOUS-WASHED. Choiseul to Guadalcanal, and Vella Lavella to Tetipari Cro-11 & 13-14 ; northern and central Fiji Dr8—2r. (25) WINGS RUSSET-EDGED. Solomons except San Cristobal Cg-16 (mixed with olive from Choiseul and Kulambangra to Florida Cro & 15) ; northern and central Fiji D18—z1 (mixed with olive on Koro D2ob). (26) TAIL PARTLY BLACK. San Cristobal C17; north-western and northern Australia, dahli and Snow Mountains E22d-23, 25 & 27; P. soror. The variable hybrid populations on the small islands off Shortland E26 have largely been omitted from the character-geography. They are discussed on p. 156. NATURAL GROUPS Co-variation and character-complexes Single characters might be used to link populations into as many sets of overlapping assemblages as there are characters under consideration. If these bore no relation to one another, or were strictly correlated with environmental features, little could be inferred about relationships and evolutionary history. In many groups, the few characters available in ordinary museum material may be too sporadic or too liable to parallel evolution to be helpful. But where there are many characters, capable of independent variation and not exclusively related to environmental differences, major discontinuities can be detected despite the local elimination or independent origin of single characters. In the P. pectoralis superspecies, both the male and the female patterns are subject to a great deal of variation. The distribution of each character is different, so that all are at least partly independent. They are of very unequal systematic value ; great when their boundaries coincide with major discontinuities in the constellation of characters, small when they occur in forms not otherwise connected. Thus the loss of the male gorget is an important character, since in other respects also the Malaitan race C12 connects with those of northern Fiji Dr8—19, and these with P. flavifrons. Hen-feathering, on the other hand, is unimportant, since the hen-feathered races A5, C16 and F33 resemble, not one another, but the females of neighbouring dimorphic forms. A character may be important in one part of the range though not in another. For example, grey-winged males distinguish the closely-related forms (group E) which range from midwestern Australia to the Bismarcks E22-25, from others whose ranges they approach closely and whose males are otherwise very similar (F31, H47-50, P. soror 3): but such grey wings are found also in eastern Fiji D19 & 21 & H56, in Tasmania and southern Australia F28—-30, on Damar H4r1 and in P. soror 2; and not in the Snow Mountain race E27, which also belongs to group E. THE PACHYCEPHALA PECTORALIS SUPERSPECIES 149 Striking variants of the male pattern, which seem to be relatively stable, mostly mark off local groups of populations with rather compact geographical boundaries. However, many of them are associated with very different character-complexes in different parts of the species range, and even locally a single character is seldom precisely co-extensive with the complex. The major discontinuities cannot be ade- quately defined by using these few more or less clear-cut characters alone. The much greater number of obviously quantitative differences must be considered as well. Although all are liable to parallel evolution and extensive intergradation, discon- tinuities are marked by concordant changes in a number of characters. Intergradation Few of the discontinuities are sharply defined. Almost everywhere, very distinct forms are connected by populations which are intermediate in range and character. Sometimes the changes are more or less regular in all the characters concerned, sometimes they are abrupt or out of step. At a few points (p. 160) the intermediate populations are highly variable, and it is clear that the intergradation is secondary— forms which differentiated in isolation have met and exchanged genes in a hybrid zone. Where the individual populations do not show exceptional variability, the intergradation may be primary or secondary. Primary intergradation could result from selection in relation to environmental gradients, or from incomplete isolation between diverging populations, or (conceivably) from the expression of orthogenetic trends after successive expansions of range (cf. Mayr & Moynihan, 1946, 1). Secondary intergradation without increased variability would imply that the hybrid populations had been stabilized by subsequent selection. In P. pectoralis the geographical patterns of intergradation, and the characters involved, make it seem highly probable that the most striking examples of intergradation have resulted from the hybridizing of differentiated forms, with subsequent stabilization (p. 160). In other areas, how- ever, character gradients seem to be correlated with climatic differences (p. 186). Subspecies—groups There are all degrees of phenotypic discontinuity, from differences between individuals of the same population upwards. The precise scope and rank of any natural group of populations must be to some extent a matter of opinion, except that sympatry introduces an objective criterion at the species level. The useful scope of the subspecies is discussed on p. 175. For convenience, such a richly diverse species as P. pectoralis must be broken up into subspecies-groups. These will not be of equal distinctiveness and homogeneity. Where a rather compact range is occupied by closely related forms, yet divided by considerable discontinuities, the number of groups to be recognized is to some extent a matter of choice. For example, the Australian subspecies-groups E and F might be combined into one, since the differences between them are much less sharp than those which mark off some other groups (cf. Mayr, 1954a, 19). Where unlike forms intergrade they should not be separated if the connecting cline seems to be environmentally determined. Mathews (1930) not only removed the 150 THE PACHYCEPHALA PECTORALIS SUPERSPECIES Northern Australian group E from P. pectoralis but split it into two species, on the basis of characters which strongly suggest selective adaptation to climate (p. 187). But where the intergradation seems to be secondary and the end-forms are sufficiently unlike, they should be separated even though the position of the dividing line will have to be arbitrarily decided. The situation in P. pectoralis is extremely complex. It seems to result from great plasticity in plumage characters combined with unusual ethological and genetic tolerance, and high mobility in successive waves of colonization combined with philopatry in local populations. Asa result, very unlike forms have evolved and then met and interbred freely, and the resulting character-gradients mimic the true adaptive clines which also occur. It is unlikely that two students will ever agree on every detail of this confused situation. In the following descriptions, the superspecies has been divided into eleven groups. These are treated in an order determined by higher grouping according to characters and trends (p. 168), regardless of the implications of sympatry and intergradation. The use of binomens for some of these groups (e.g. “ P. schlegelii’’) and informal designations for others (e.g. “‘ Solomons group C’’) anticipates the findings of later sections. A description of the divergence of each group as a whole from the standard patterns is followed by character-geographies of departures from the group patterns so determined. In general, group patterns have been arrived at (like the standard patterns of the whole superspecies) by combining those characters which are found in a majority of the contained forms, excluding those which are obviously intermediate with other groups. But in the Lesser Sunda Isles, Moluccas and Fiji, it seems clear that few forms have escaped contamination by gene-exchange. Here the group patterns are taken to be those of the apparently pure forms Ar, Bo-7, and D18a & Iga. P. schlegelii (Text-fig. 3, p. 167) Small birds. Male chin black, throat-patch small; gorget very broad, fore-edge defined by white feathers; breast, flanks and belly rufous-washed ; mantle very dark ; wing, upper tail-coverts and tail black. Female throat barred grey and white, gorget grey, breast and flanks olive ; belly lemon-yellow ; cap and auriculars grey, mantle pure olive. MALE VaRIATION. Throat-patch larger and gorget narrower in Vogelkop I. Rufous very deep in south-eastern New Guinea 3b, paling steadily westwards to 3a ; much paler in Vogelkop and Cyclops Mountains 1-2. FEMALE VARIATION. Throat white with grey bars, gorget and breast pale in Vogelkop and Cyclops Mountains 1-2; throat grey with white bars, gorget and breast dark from Weyland Mountains to south-eastern New Guinea 3. Greys and olives brownish in Vogelkop 1. Lesser Sundan group A (Text-fig. 2, p. 161) Very small birds. Male chin black, throat-patch small and separated by black feathers from auriculars; gorget narrow, fore-edge defined by white feathers ; breast, belly and flanks rufous-washed ; collar narrow and olive-washed ; upper —— THE PACHYCEPHALA PECTORALIS SUPERSPECIES 151 tail-coverts and tail black. Female throat whitish ; gorget greyish, pale and vague ; cap sandy-grey ; mantle pale sandy-olive. MALE VARIATION. Hen-feathered on Salayer A5—male differs from female in larger throat-patch, greyer gorget and cap, grey auriculars, and darker and greener mantle. Throat-patch touches auriculars from Java to Alor and in Flores Sea A2-4; patch not small, and fore-edge of gorget with black tips, in Flores Sea A4. Rufous pale on belly in Java Aza, more so on Bali A2b, restricted to band behind gorget from Sumbawa to Alor A3, very pale and barely detectable below breast in Flores Sea A4. Wing black and mantle mottled with black in Flores Sea A4 (individually variable). Tail with narrow pale tips from Java to Alor and in Flores Sea A2-4. FEMALE VARIATION. Throat pure white in Flores Sea and on Salayer A4-5, pinkish-buff from Java to Alor A2-3. Gorget rather pinkish from Java to Alor and on Salayer A2-3 & 5; breast and belly uniformly pinkish in Flores Sea Aq. Breast and belly with very little yellow from Java to Alor and in Flores Sea A2-4. Cap almost pure grey and mantle pure olive on Salayer A5, distinctly sandy from Sumbawa to Alor and in Flores Sea A3-4, less so on Sumba and Java AI-2. Except for their smaller size, females from the Flores Sea A4 are very like those of Timor H39. Moluccan group B (Text-fig. 2) Medium-sized to large birds. Male chin black; throat feathers long, partly covering gorget and cutting it off from auriculars ; gorget narrow from side to side, without black tips in the fore-edge ; upper tail-coverts and tail black. Female throat barred grey and white, gorget grey, breast and flanks pale olive ; belly lemon- yellow ; cap and auriculars grey, mantle pure olive. SIZE VARIATION. Large birds on Ternate, Tidore and Obi B7-8. MALE VARIATION. Chin white, throat feathers shorter, and tail with narrow and obscure pale tips on Obi B8. FEMALE VARIATION. Throat-bars and gorget buff, breast and flanks ochraceous, belly golden-yellow, cap and mantle brownish on Obi B8. Greys slightly paler and olives yellower on Ternate and Tidore B7 than from Morotai to Batjan B6. The Obi form B8 is precisely intermediate between those of Ternate and Tidore B7 and Ceram H43, except that the female belly is deeper yellow than in either. Solomons group C (Text-fig. 2) Very large birds. Male chin black; throat-patch yellow, rather small; gorget broad, fore-edge defined by yellow feathers ; collar washed with olive. Female bill pale ; gorget broad and vague ; breast and flanks washed with melanins ; cap olive ; wing-feathers edged with russet. SIZE VARIATION. Small birds on Rennell and San Cristobal C16—17, large (not very large) from Vella Lavella to Gatukai C13 & 15. 152 THE PACHYCEPHALA PECTORALIS SUPERSPECIES MALE VARIATION. Hen-feathered on Rennell C16—male almost indistinguishable from female, but with slightly more carotenoid on the average. Chin yellow on San Cristobal C17. Throat-feathers long (extending behind auriculars), gorget very broad on Vella Lavella and Ganonga C13. Gorget absent, but throat-feathers sometimes black-fringed, on Malaita C12. Some black tips in fore-edge of gorget on San Cristobal C17. Rufous patch below gorget on Santa Anna Cr7b and in individuals on San Cristobal Ci7a. Head olive with blackish lores and auriculars on Santa Anna C17b, varies from olive (auriculars brownish) to black on San Cristobal C17a. Collar absent on Vella Lavella and Ganonga C13; reduced to a trace on Malaita and from Kulambangra to Gatukai and Tetipari C12 & 14-15 ; reduced to a trace on the hind-neck, but with broad patches laterally, on San Cristobal C17. Entire upperparts and flanks black on Vella Lavella and Ganonga C13 (individuals on Ganonga C13b have narrow olive edges on the wings) ; wing black and mantle more or less mottled with black on Rendova and Tetipari C14 ; centres of mantle- feathers blackish, and olive dark greenish, from Kulambangra to Gatukai C15 ; olive rather greenish on San Cristobal C17. Upper tail-coverts olive from Kulambangra to Gatukai and on San Cristobal C15 & 17; olive edges broad on Malaita C12. Tail more or less olive on San Cristobal Cr7. FEMALE VARIATION. Bill straw-coloured from Kulambangra to Guadalcanal and Rennell Croc-11 & 15-16 ; blackish brown elsewhere. Individuals on Vella Lavella C13a are melanistic, with black gorgets and more or less black upperparts. Underparts faintly washed with yellow, cheeks yellower than throat, on Bougain- ville and Guadalcanal (juveniles sometimes much yellower) and Malaita Cg, 11 & 12 ; yellow pale on Rendova, Tetipari and Rennell C14 & 16; lemon-yellow elsewhere (when unmixed with melanins). Throat faintly barred on Malaita C12; underparts (except mid-belly) conspicuously shaft-streaked on Malaita C12, more faintly on Bougainville, Choiseul and Guadal- canal, and from Kulambangra to Gatukai Cg—10a, 11 & 15. Gorget greyish on Bougainville and Malaita Cg & 12; olive from Choiseul and Kulambangra to Florida, and on Rennell and San Cristobal Cro & 15-17 (fairly narrow and distinct on San Cristobal C17) ; rufous on Guadalcanal and from Vella Lavella to Tetipari Crz & 13-14. Underparts washed with deep rufous on Rendova and Tetipari C14 ; rufous very variable in extent and depth on Vella Lavella and Ganonga C13 ; confined to gorget, breast and flanks on Ysabel and Guadalcanal Crob & 11 (deep, pale or absent), and Choiseul, Russel Isles and Rennell Croa & c & 16 (pale). Breast and flanks pale olive from Kulambangra to Gatukai and on Rennell C15-16, and in individuals from Choiseul to Florida Cro. Cap citrine, mantle olive on San Cristobal C17; cap brownish or rufous olive, mantle olive from Kulambangra to Gatukai and on Rennell C15-16 ; cap brownish THE PACHYCEPHALA PECTORALIS SUPERSPECIES 153 citrine, mantle citrine from Buka to Malaita Cg-10 & 12 (sometimes with a rufous wash on Ysabel Crob, less common on Choiseul Croa) ; cap rufous, scapulars and mantle strongly washed with rufous (most consistently on Rendova and Tetipari C14) on Guadalcanal and from Vella Lavella to Tetipari Crz & 13-14. Indications of a collar from Buka to Guadalcanal and on Vella Lavella and Ganonga Cg-11 & 13. Olive dark from Kulambangra to Gatukai C15, darker still on Rennell Cx16. Olive dull (little carotenoid) on Rennell C16 ; richer on Guadalcanal, Rendova and Tetipari C11 & 14; richer on Bougainville and from Kulambangra to Gatukai Co & 15; richer from Choiseul to Malaita and on Vella Lavella and Ganonga Cro, 12 & 13; rich on San Cristobal C17. Wing-feathers edged with olive on San Cristobal C17; russet mixed with olive from Russel Isles to Kulambangra Croc & 15, less strongly from Choiseul to Florida Croa-b. Tail pure olive, with variable black subterminal patch, on San Cristobal C17 ; citrine from Buka to the central Solomons and Guadalcanal Co, 13-15 & 10c-I1; olivaceous brown from Choiseul to Malaita Croa—b & 12 ; dull brown on Rennell C16. Fijian group D (Text-fig. 2) Medium-sized birds. Male chin and throat yellow ; no gorget ; forehead yellow ; black of cap continued on to hind-neck, collar reduced to a trace ; Olive very dark ; upper wing-coverts yellow-edged ; upper tail-coverts olive ; olive tips of tail-feathers dull but wide. Female forehead, circumoculars, cheeks and under tail-coverts faintly washed with yellow, underparts otherwise without carotenoid (buffy-grey, with throat and mid-belly slightly paler) ; underparts streaked with brown, and throat and breast barred also ; upperparts dull olive-brown, wing-feathers edged with dark Tusset ; individuals are dark rufous all over. MALE VARIATION. Underparts lemon-yellow on Viti Levu and Ovalau D18, orange-yellow elsewhere. More or less complete gorget, individually variable, on Ovalau and from south- western Vanua Levu to Kio D18b & 1gb-c; gorget complete (though somewhat irregular) on Taviuni, Koro and Vatu vara D20-21. Breast, belly and under tail-coverts washed with brown on Vatu vara Dat ; under tail-coverts ochraceous on Koro D2ob, decreasingly so towards Vanua Levu D20a to Igb. Collar narrow but uninterrupted on Vatu vara Dat. Wing-quills edged with greyish on Vanua Levu and Vatu vara Dig & 21, becoming more olive eastwards, towards Viti Levu and Koro D18 & 20. Upper tail-coverts with black centres on Koro and (larger) on Vatu vara D2ob-21. FEMALE VARIATION. Bill brown on Rambi and Kio Drgc. Underparts evenly-coloured on Koro and Vatu vara D2ob & 21 ; mottling becomes more common and emphatic towards the large islands D18b to a, and 20a to Iga. Ground-colour of underparts cinnamon on Koro D2ob and (rather paler) Vatu vara D21, individually variable, becoming greyer, towards the large islands D18b to a, and 20a to Iga. Forehead, circumoculars and cheeks not yellow on Koro and Vatu vara Dzob-21 ; increasingly so towards the large islands D18b to a, and 20a to 1ga. 154 THE PACHYCEPHALA PECTORALIS SUPERSPECIES Females on Koro and Vatu vara D2ob-z1 are very like those in southern Fiji H54-56. P. flavifrons (Text-fig. 5, p. 173) Small birds. Male chin dull black, throat dull black with broad yellow or white tips; no gorget ; underparts lemon yellow ; forehead yellow or white ; upperparts dull black (slightly washed with olive), primaries with obscure narrow greyish edges. Female cock-feathered—like the male, but throat pale grey with narrow yellow or white tips, forehead blackish (with or without a yellow wash). INDIVIDUAL VARIATION. The tips of the throat feathers are usually yellow, but may be white ; two specimens in the British Museum (Natural History) have mixed yellow and white tips. Some white-throated birds have the forehead white (in females, without a yellow wash). The distribution of these phases in the collections of the British Museum (Natural History), and the American Museum of Natural History (from Mayr, 19326), is shown in Table II : there is no evidence of a significant difference in representation of the colour phases, either between the sexes or between Savaii and Upolu. All the B.M. (N.H.) specimens are from Upolu. TaBLE II.—Colour Phases in P. flavifrons. Forehead. a SS Throat. Yellow. White. Yellow . é . 3 48 — Yellow and white 2 — White 8 9 P. soror (Text-fig. 3) Small birds. Male throat-patch rather small; gorget narrow ; underparts lemon- yellow ; breast and flanks faintly olivaceous ; collar reduced to a trace; mantle very dark greenish olive; edges of wing quills greyish ; upper tail-coverts olive, tail partly olive. Female throat-patch white and small; gorget narrow, clear and brownish ; breast and flanks pale citrine, yellow wash extending high on breast ; belly pale lemon-yellow ; cap olivaceous-brown ; tail partly black. MALE VARIATION. Mantle increasingly golden-olive towards the west 3 to I. Wing quills greyest in the centre 2. Tail olive, with a variable black subterminal patch, in the west and east 1 & 3; black, with wide olive edges basally, in the centre 2. FEMALE VARIATION. Upperparts much browner in Vogelkop 1; increasingly purer olive towards the east 2 to 3. Northern Australian group E Very small to medium-sized birds. Male throat-feathers white to their bases ; edges of wing-quills grey ; edges of upper wing-coverts yellow ; primary-coverts black. Female throat white, with barring ; gorget well defined, buffy-grey ; breast, e THE PACHYCEPHALA PECTORALIS SUPERSPECIES 155 flanks and belly whitish to golden-yellow, without melanins; cap rather pure grey, mantle pure olive (or grey) ; tail partly black. SIZE VARIATION. Verysmall birds in mid-western Australia E22a-c, size increasing through north-western Australia E22d to small in northern Australia and southern New Guinea E23-24. Like the Snow Mountain birds E27, those grouped together as dahli E25 are mostly medium-sized—but there is considerable variation between the scattered populations, with the birds of Fergusson E25c (and Teste and Long Islands E25b & d) rather large, and those of south-eastern New Guinea and Witu E25a & e apparently rather small. MALE VARIATION. Throat-feathers grey-based in Snow Mountains E27 (Text-fig. 3). Breast and belly lemon-yellow in mid-western Australia E22a, gradually becoming deeper northwards E22b to d, golden-yellow from northern Australia eastwards E23-27. Collar narrow in midwestern Australia E22a (widening northwards Ez22b to 23), and Snow Mountains E27. Mantle greenish-olive in midwestern Australia E22a (becoming yellower north- wards E22bto 23), and Snow Mountains E27. Edges of wing-quills olive in Snow Mountains E27. Primary-coverts olive in mid-western and north-western Australia and Snow Mountains E22 & 27 ; sometimes with very narrow pale edges from northern Australia to southern New Guinea E23 to 24. Upper tail-coverts olive in mid-western and north-western Australia E22. Tail with broad olive edges basally in midwestern Australia E22a, decreasing northwards Ez22b to d. FEMALE VARIATION. Females from Teste and Fergusson E25b-c are unknown. Throat unbarred in mid-western Australia E22a-c, barring pale in north-western and northern Australia E22d—23 ; throat and gorget shaft-streaked (in addition to barring) from Cape York to southern New Guinea E24 (faintly) and in dahli and Snow Mountains E25 & 27. Breast and flanks pale olivaceous in Snow Mountains E27 ; ochraceous from Cape York to southern New Guinea E24. Breast and belly whitish and under tail-coverts pale yellow in mid-western Australia E22a-c ; breast and belly pale yellow, under tail-coverts lemon-yellow, in north- western Australia, and from Cape York to southern New Guinea E22d & 24 ; under- parts lemon-yellow in northern Australia E23, golden-yellow in dahli and Snow Mountains E25 & 27. Cap brown in Snow Mountains E27. Mantle grey, only rump olivaceous, in mid-western Australia E22a—c, becoming olive northwards E22d to 23. Olive bright in Snow Mountains E27. Edges of wing feathers, rump and base of tail brownish on Witu, Malie and Nissan E25e & j-k. Tail olive in mid-western Australia, and from Cape York to southern New Guinea E22a-c & 24; with a variable amount of black in north-western and northern Australia E22d-23 ; more than half black in dahli E25; black with olive edges in Snow Mountains E27. Mayr has shown (1932a) that the birds on small islands west of Shortland E26 156 THE PACHYCEPHALA PECTORALIS SUPERSPECIES form a hybrid population. The nine known males show almost every degree of inter- mediacy between dahli E25 and the Bougainville-Shortland race Cg—except that they do not have black chins like those of the Solomons group C. In this series the largest birds have the yellowest throats, and the smallest ones the whitest. Other characters seem to vary independently, so far as can be judged from this short series. There is rather less variation between the nine females, which are also inter- mediate between those of E25 and Cg. The females from Cape York to southern New Guinea E24 differ from those on either side (E23 & 25) in characters which approach those of the Queensland form F31—breast and flanks ochraceous, little ventral yellow, tail without black (Mayr, 1954a). The Louisiades form H47 is intermediate between groups E and H. It is here placed in the latter because the male has olive edges to the wings, and the female is brownish ; but the white throat, sharp and narrow gorget and bright yellow under- parts of the female are untypical of group H. The Tabar form H50 may be similarly intermediate (p. 164). Southern Australian group F Large birds, with long wings and tails, small bills and soft plumage. Male breast and belly lemon-yellow ; collar rather narrow and washed with olive ; upper wing- coverts yellow-edged ; upper tail-coverts olive, base of tail olive or grey. Female throat barred ; gorget, breast and flanks buffy-grey, belly more or less washed with melanins ; yellow absent from underparts, or only in under tail-coverts ; cap grey, mantle dull sandy-olive. SIZE VARIATION. Large birds with long tails and very small bills in Tasmania F2g, size and relative tail-length decreasing and bills lengthening northwards— smallest birds near Cairns F31b, longest bills on Lord Howe and Norfolk Islands F32-33. MALE VARIATION. Hen-feathered on Norfolk Island F33—indistinguishable from female. Yellow somewhat paler from south-western Australia to Tasmania F28—29. Collar broader and clearer on Lord Howe F32. Wing quills blackish brown with grey edges in Tasmania F29, centres blackening and edges becoming more olive northwards F28 and 30 to 31. Tail without black in Tasmania F29 ; about one-third black from south-western Australia to South Australia F28a—b; increasing through Victorian mallee F28c to two-thirds black in south-eastern Australia F30 ; two-thirds black on Lord Howe F32 (olive tips very wide) ; mainly black, olive varying from wide edges to a wash at the extreme base, in Queensland F31 (averaging blacker near Cairns F31b). Pale part of tail grey in Tasmania and southern Australia F28-30a ; olive else- where, but often greyish near Cairns F3rb. FEMALE VARIATION. Belly pinkish from south-western Australia to South Australia F28a—b, fading to buffy in Victorian mallee, Tasmania and near Cairns F28c-29 & 31b; whitish in south-eastern Australia F30 ; often faintly yellowish in THE PACHYCEPHALA PECTORALIS SUPERSPECIES 157 southern Queensland F3r1a, and always on Lord Howe F32; distinctly yellow on Norfolk Island F33. Under tail-coverts whitish in Tasmania and southern Australia F28—30. Upperparts rather brownish in Tasmania F29, and more or less brownish in most individuals near Cairns F31b ; rather pure grey in south-western Australia F28a. Mantle grey in Tasmania F29; sometimes with a faint olive wash in southern Australia F28 & 30; dull olive elsewhere. Tail grey in Tasmania and southern Australia F28-30; dull olive near Cairns F31b ; olive elsewhere. Southern Melanesian group G Small birds. Male gorget narrow ; breast and belly lemon-yellow ; collar narrow and washed with olive ; centres of wing-feathers blackish-brown ; upper tail-coverts and tail olive. Female throat white; gorget greyish; breast and belly without melanins. SIZE VARIATION. Very large birds on Loyalty Islands G35. MALE VARIATION. Lateral throat feathers long, separating gorget from auriculars, on New Caledonia G34 (Text-fig. 5). Gorget dull black, and very narrow, on New Caledonia G34 ; dull black on Aneitum G36 ; rather broad on Vanikoro G38. Pale rufous patch below gorget on New Caledonia G34. Flanks and under tail-coverts ochraceous, rump, tail and edges of secondaries washed with brownish, in individuals on Aneitum G36. Breast and belly golden-yellow on Loyalty Islands G35. Cap and auriculars grey on New Caledonia G34; dull black, cap more or less scalloped with olive, on Aneitum G36. Collar reduced to a trace on New Caledonia G34, very narrow on Erromango G37a. Mantle greenish-olive on Erromango, and from Raga to Vanikoro G37a & 37e-38 ; golden-olive on Loyalty Islands, and Malekula and Santo G35 & 37d. Upper tail-coverts black-centred, tail black, on Vanikoro G38; tail with black patches (on inner webs of all but central quills) from Erromango to Banks Islands G37. FEMALE VARIATION. Throat and gorget shaft-streaked on Vanikoro G38. Gorget olive on Loyalty Islands and Vanikoro G35 & 38 ; greyest on Erromango, Efate and Banks Islands G37a-b & f; brownish on Aneitum G36. Gorget extends into buffy-grey wash on breast and belly on New Caledonia G34. Underparts golden-yellow on Loyalty Islands G35, lemon-yellow on Vanikoro G38 ; elsewhere, under tail-coverts pale yellow, breast and belly faintly washed with yellow (palest on Malekula and Santo G37d). Cap and auriculars grey on Vanikoro G38; cap greyish on Efate and Banks Islands G37b & f; brownish on Aneitum G36. Mantle pure golden-olive on Loyalty Islands G35, pure greenish-olive on Vanikoro G38 ; elsewhere dull olive, purest on Efate and Banks Islands G37b & f, brownish on Malekula and Santo G37d ; very brown on Aneitum G36. 158 THE PACHYCEPHALA PECTORALIS SUPERSPECIES Tail and edges of secondaries browner than mantle on Loyalty Islands G35, some- what so on Aneitum and Erromango G36-37a. JUVENILE VARIATION. Throat lemon-yellow on Vanikoro G38, sometimes yellow- washed in New Hebrides (recorded from G37c & f by Mayr, 19325). Juveniles on Vanikoro G38 are remarkably like those on San Cristobal C17, except for the shaft- streaking of their throats and gorgets. The New Hebrides G36-37 are a region of incipient subspeciation, with the females of almost every island distinguishable in series (Mayr, 19326). The Aneitum popula- tion G36 is rather distinct, and that on Erromango G37a more so than the others. Widespread group H (close to standard patterns, Text-fig. 1) Small to very large birds. Male of standard pattern. Female underparts rather uniform, brownish, with little yellow; upperparts brownish, cap not contrasting, edges of wing feathers distinctly browner than mantle. SIZE VARIATION. Very large birds on Tenimber Isles, Lihir and Tonga H42, 49 & 57; large on Tabar and Lau archipelago H50 & 56; small on Timor and Damar, Sula and Peleng Isles, large islands in the Bismarcks, and Kandavu H309, 41, 45-46, 48 & 54. MALE VARIATION. Chin and throat black in Tonga H57 (Text-fig. 5) ; yellow on Tenimber Isles H42. Throat-feathers long, cutting gorget off from auriculars (fore-edge of gorget defined by white feathers) on Ceram, and Ndeni and Santa Cruz islets H43 & 52; gorget very narrow near auriculars, with few black tips, on Buru H44. Gorget streaked with white on Babar H4o ; dull black and very narrow on Kandavu H54, narrow on Ngau H55. Underparts lemon-yellow from Timor to Tenimber Isles and on Utupua, Kandavu and Tonga H39-42, 53-54 & 57 ; orange-yellow in Louisiades and on Lihir and Manus H47, 49 & 51. Under tail-coverts ochraceous on Ngau H55. Upperparts (except for narrow olive edges on wing-feathers) black on Ndeni and Santa Cruz islets H52 ; mantle mottled with black in Lau archipelago H56, somewhat so on Ngau H55. Primary-coverts black on Ceram and Buru, Ndeni and Santa Cruz islets, and Lau archipelago H43-44, 52 & 56. Collar absent on Ndeni and Santa Cruz islets H52 ; reduced to a trace on Tenimber Isles H42, and Ngau and Lau archipelago H55-56 (black of cap extends on to hind- neck) ; washed with olive from Timor to Damar, in the Louisiades and on Utupua and Kandavu H39-41, 47 & 53-54. Olive dark and greenish on Buru, Ngau and Lau archipelago H44 & 55-56; dull on Kandavu H54. Olive washed with brownish (especially. on rump, tail and edges of secondaries) on Ceram and Louisiades H43 & 47. Edges of wing quills grey on Damar H4r1, and greyish in Lau archipelago H56. Upper tail-coverts olive on Timor, Tenimber Isles, Louisiades, Kandavu and Tonga THE PACHYCEPHALA PECTORALIS SUPERSPECIES 159 H39, 42, 47, 54 & 57; black from Ceram to Peleng Isles H43-46 (very narrow olive edges on Buru H44). Tail olive on Timor and western Louisiades H39 & 47a; olive with blackish subterminal patches on Misima and Kandavu H47b & 54; olive basally on Babar and Rossel H40 & 47c ; with broad olive edges basally on Damar H4t. Pale tip of tail very broad and yellowish in Tonga H57. FEMALE VARIATION. Throat white on Timor and Babar, Louisiades and Utupua H39-40, 47 & 53; whitish on Damar, Ceram, Sula and Peleng Isles, Tabar, Manus, Ndeni and Santa Cruz islets H41, 43, 45-46 & 50-52. Throat washed with yellow on Tenimber Isles H42. Throat barred on Damar, Sula and Peleng Isles H41 & 45-46, and faintly in Tonga H57. Throat and gorget streaked on Manus H51, and faintly on Tabar H50. Underparts uniformly cinnamon in southern Fiji H54—-56 (very richly so on Ngau H55). Gorget scarcely darker than throat and belly on Mussau and Lihir H48d—4o ; little darker on Tenimber Isles, large islands of the Bismarcks, and Ndeni and Santa Cruz islets H42, 48a—c & 52. Gorget little darker than breast, but greyer, on Timor and Damar and from Ceram to Peleng Isles H39, 41 & 43-46. Gorget pale and narrow on Babar and Louisiades H4o & 47 (vinous), Tabar, Manus and Tonga H50-51 & 57 (cinnamon) ; broad on Utupua H53 (dark brown). Breast and belly without melanins in Louisiades H47; belly without melanins, breast faintly cinnamon on Tabar and Tonga H50 & 57; belly without melanins, breast and flanks brownish on Utupua H53; breast and belly faintly washed with cinnamon on Babar and Manus H4o & 51. Breast and belly without yellow on Ngau and Lau archipelago H55-56; faintly washed with yellow on Timor, Damar, Buru and Kandavu H309, 41, 44 & 54, more strongly on Tenimber Isles, Ceram and Sula Isles H42—43 & 45 ; yellow pale on Babar, Bismarcks except Tabar, and Ndeni and Santa Cruz islets H40, 48-49 & 52; deep in Peleng Isles and Louisiades H46 & 47, and especially on Tabar and Manus H50-51. Cap grey on Tabar, Ngau and Tonga H50, 55 & 57; distinctly greyer than mantle on Manus, Utupua, Kandavu and Lau archipelago H51, 53, 54 & 56; distinctly browner than mantle on Bismarcks except Tabar H48—49. Mantle bright citrine on Tabar, Manus and Tonga H50-51 & 57; dull sandy- olive on Timor, Damar, Tenimber Isles, Kandavu and Lau archipelago H39, 41-42, 54 & 56; brownish olive on Babar H40 (tail pure olive), Peleng Isles, Louisiades, large islands of the Bismarcks, Utupua and Ngau H46, 47, 48a-c, 53 & 55; browner and duller from Ceram to Sula Isles and on Ndeni and Santa Cruz islets H43-45 & 52; olive-brown on Mussau and Lihir H48d—49. Edges of wing feathers scarcely browner than mantle on Tabar, Manus and Tonga H50-51 & 57. Not only are the distinctive characters of the group few and slight, but most of the forms are intermediate in some respects with neighbouring groups. Besides penetra- tion by characters proper to other groups, intermediate forms seem to be liable to special changes (p. 165). The wide range of this group has enabled it to meet and 160 THE PACHYCEPHALA PECTORALIS SUPERSPECIES intergrade with five of the seven other subspecies-groups, and there seems to have been gene-flow into it in the following areas : From group A to Timor and Babar H39-40. From group B to Ceram, Peleng and Sula Isles H43 & 46-45. From group D to southern Fiji H54—56. From group E to Louisiades H47 (and possibly Damar, Tabar and Manus H4r & 50-51). From group G to Utupua H53. When the effects of this gene-exchange have been allowed for, three forms stand out conspicuously from the remarkably uniform remainder : Tenimber Isles H42 (very large, throat yellow). Ndeni and Santa Cruz islets H52 (male upperparts black, throat-feathers long). Tonga H57 (very large, male throat black, female with little brown and much yellow). INTERGRADATION AND BRIDGELESS GAPS Gene-exchange between subspecies-groups The eight subspecies-groups of P. pectoralis approach one another in several areas, in all of which there are signs of intergradation between them (Text-fig. 8). Groups A and H intergrade in the Lesser Sunda Isles, B and H in the southern Moluccas, C and E in the western Solomons, C and G in the eastern Solomons, D and H in central Fiji, E and F in southern New Guinea, E and H in the Louisiades, and G and H in the Santa Cruz archipelago. In most of these areas, it is not immediately obvious whether the intergradation is primary or secondary. Only in the western Solomons is there indisputable evidence of hybridization between very dissimilar forms. The populations on three islets west of Shortland (E26) are highly variable. Extreme individuals closely resemble the Shortland race Cg on the one hand and dahli E25 on the other, while every degree of intermediacy is found (p. 156). On San Cristobal Cr7a also there is considerable variation between males, in the amount of rufous on the breast and of black on the head and tail. Their song, too, is remarkably variable (Cain & Galbraith, 1956). Though the San Cristobal race agrees with the rest of group C in the yellow throat, and the olive cap and pale bill of the female, it differs from the standard pattern of that group in most other characters. It is much smaller. The male has a yellow chin, black tips in the fore-edge of the gorget, a rufous breast-patch, and partly olive head and tail. The female is bright yellow underneath, with a narrow gorget and no melanic wash, lacks rufous on the upperparts (including the wings), and has a partly black tail. Many of these characters of the males are retarded, and of the females advanced. Avifaunally San Cristobal is distinct from the rest of the Solomons, with several endemic forms showing markedly reduced size (Collocalia esculenta makirensis, Ptilinopus solomonensis solomonensis, Halcyon chloris solomonis, Rhipidura rufifrons russata, Monarcha vidua, Myiagra cervinicauda and Aplonis grandis dichrous), reduced sexual dimorphism (Coracina tenuirostris salamonis, C. lineata makirae, THE PACHYCEPHALA PECTORALIS SUPERSPECIES TE Fic. 2.—Four extreme forms of P. pectoralis (schlegelit assemblage) which intergrade with more nearly standard ones (sovor assemblage). Sumba Ar; Ternate B7; Shortland Cg; Vanua Levu Diga. 161 162 THE PACHYCEPHALA PECTORALIS SUPERSPECIES Myiagra cervinicauda, Myzomela nignita tristrami and Dicaeum tristramt), or increased variability (Ptilinopus solomonensis solomonensis, Halcyon chloris solomonis, Coracina lineata makivae and Petroica multicolor polymorpha) in comparison with their western representatives. These trends may result simply from the isolated peripheral position and impoverished avifauna of the island. However, several birds on San Cristobal and its outlying islands are derived from southern Melanesia rather than from the rest of the Solomons (Ptilinopus richards, Lalage leucopygia, Vitia parens, Rhipidura fuliginosa, Myiagra cervinicauda and Myzomela cardinalis). The vari- ability of the male of P. pectoralis here suggests the possibility of hybridization, which might have disturbed the sexual dimorphism (p. 165). The Vanikoro race G38 shows some of the characters to be expected in the other putative parent : the female is rather similar to that on San Cristobal, except for its streaked white throat, while the juvenile has a streaked but yellow throat. The San Cristobal race shares with most members of group G the pale tail of the male and the melanin-free throat of the female ; and with the New Caledonian race G34 the rufous breast-patch and pale cap. In the light of extensive hybridization between unlike forms elsewhere, it seems most probable that the San Cristobal race is a hybrid between groups C and G. The New Caledonian race G34 differs from its neighbours of group G in several systematically important characters of the male plumage, which suggest affinities with P. schlegelii and groups A to C of P. pectoralis (the schlegelii assemblage, p. 168). The elongation of the white feathers at the sides of the throat and the failure of the gorget to join the auriculars are reminiscent of group B. The absence of black tips in the fore-edge of the gorget (laterally) is characteristic of the whole assemblage. The rufous breast is found elsewhere only in P. schlegelii and group A, and on San Cristobal, and is in the form of a patch rather than a band only on New Caledonia and San Cristobal. These two forms share also the pale cap, and the suppression of the collar (except for lateral patches in the latter). Though the females on these two islands are very different, that on New Caledonia agrees with typical females of group C rather than of group G in having the belly washed with melanins. The number of characters involved makes it improbable that they have been developed independ- ently, and it seems likely that this race too has had genetic contributions from group C as well as group G. Two further forms, here assigned to group H, show characters which are elsewhere systematically important and confined to the schlegelii assemblage (see Table III). The race on the Tenimber Isles H42 has a yellow throat. That on Ndeni and the Santa Cruz islets H52 has overlapping throat feathers without black tips, and a black mantle. But in other respects these forms agree well with members of group H. It would not be justifiable (without further evidence) to deny the possibility of inden- pendent origin for these characters, and to postulate mixed ancestry for these forms also. A latent tendency towards yellowing of the throat is apparent in many forms (p. 142). In the melanic race H52 the elongation of the throat feathers and the exclusion of melanin from their tips may be significant in maintaining the con- spicuousness of the throat-patch despite encroachment by black areas (as in the other melanic race C13). THE PACHYCEPHALA PECTORALIS SUPERSPECIES 163 The populations between Koro and the large Fijian islands (D18b & 19b-20a) are evidently hybrid (Mayr, 1932b). There is much variability—especially on Ovalau, Rambi and Kio D18b & 19c—in the degree of development of the male gorget and forehead-spots, and in the colour and streakiness of the female underparts. The average character of the populations changes progressively from Koro to the main- land of Vanua Levu (D2ob to Iga), while the population on Ovalau D18b is intermediate in character between those of Koro D2ob and Vitu Levu D18a. Unfortunately, my material is inadequate to demonstrate quantitatively the changes in hybrid index and in variability which are qualitatively apparent in Mayr’s description. The races on Koro and Vatu vara D2ob-21 are themselves intermediate in character between the extreme Fijian forms (D18a & Iga) and members of group H, to which they are linked by the races in southern Fiji H54-56. Males on Koro and Vatu vara have yellow throats with gorgets, those in southern Fiji white throats with gorgets ; the yellow forehead reappears on Vatu vara ; while all Fijian males have the collar obscure or very narrow. Females in southern and central Fiji (D2o-21 & H54-56) are all much alike—differing from those of northern Fiji in having unstreaked cinnamon underparts, and from most of those in group H in having them almost uni- form from chin to vent and with scarcely any carotenoid. Blackness of the mantle (D20 & H55-56) and greyness of the wing quills (Dig & 21 & H56) are common to forms on either side of the intergroup boundary. Group H intergrades with groups A and B also. The male on Sumba Ar has a black chin, a small throat-patch, no black tips in the fore-edge of the gorget, a rufous ventral wash and a wholly black tail. These characters are reduced or absent elsewhere in group A, the dilution being greatest in the Flores Sea Aq and least in Java Aza. Although there is a sharp change in size at the Ombai Strait (along which the boundary between groups A and H has been drawn), and the special male characters of group A do not appear east of it, the female on Timor H39 is very like that in the Flores Sea (which differs from other females of group A in having a pure white throat and pinkish belly), and there is a fairly complete series of forms leading to a typical member of group H on Buru (H39 to 41 & 44). Males in the northern Moluccas B6-7 have black chins, long throat-feathers partly covering the gorgets, and wholly black tails; while the females are unusual in their heavily-barred throats, pure greys and olives, olive breasts and grey auriculars. There is a perfect series of intermediates leading from these through Obi B8 and Ceram H43 to Buru H44. Although the male characters of group B do not appear there, the female in the Peleng Isles H46 is very like that on Obi B8, and the Sula Isles female H45 is intermediate between those of Peleng and Buru. The changes involved are quite different in these three areas of intergradation. They involve oddities of pattern, as well as the quantitative changes which seem more likely to be subject to environmental selection. Only in the Lesser Sunda Isles is there a marked climatic gradient (of increasing aridity eastwards from Java to Timor) with which the progressive change in character might be correlated. But here the character-progression (from A1—za-2b-3-4—H39) does not run parallel to that of climate. In the Moluccas the changes span only seven degrees of latitude across the equator, and in Fiji four degrees within the tropics, so that regular climatic ZOOL. 4, 4. 12 164 THE PACHYCEPHALA PECTORALIS SUPERSPECIES changes cannot be great. It seems unlikely that adaptation to environmental gradients is involved in these character-progressions, although adaptive clines might be developed in relation to subtle differences (perhaps in the fauna or flora). The geographical patterns of character-dilution suggest introgression rather than continued gene-exchange between gradually diverging populations (see Text-fig. 8). Finally, groups A, B and D seem (p. 168) to be more closely related to one another than to group H, with which they all intergrade. There is thus no reason to believe that the intergradation in these areas is primary. The situation can be explained with the maximum economy of hypothesis in terms of a recent burst of range expansion by group H, which has come into secondary contact with diverse and anciently-isolated forms and freely interbred with them. The same thing has almost certainly happened in the western Solomons, where dahl E25 is a relatively very recent arrival and the hybrid population E26 is still exceedingly variable. Meise (1936) and Mayr (1942) have suggested that the Pachy- cephala on Koro D2ob is of hybrid origin and has been genetically stabilized by subsequent selection. This seems much the most probable explanation for the other intermediate, though not especially variable, forms just considered. Since the characters which distinguish groups E to H from one another are relatively slight and mainly quantitative, it is more difficult to decide whether the intergradation between these groups is primary or secondary. On the other hand there is less reason to doubt that the rather similar forms involved can interbreed, and the character-geography strongly suggests gene-flow between these groups in several areas (Text-fig. 8). Mayr (1954a) has pointed out that, while the male in southern New Guinea E24 is like its relatives E23 and 25, the female shows (in its buffy belly with little yellow, and lack of black in the tail) dilution by characters of group F. The dilution appears to decrease into south-eastern New Guinea E26a (Rand, 1940). It is clear that gene-flow from northern Queensland F3rb is involved. Mayr also suggests that the character gradient in mid-western Australia (E22d to a) is the result of gene-flow from south-western Australia F28a. The partly olive tail of the male and the reduc- tion of carotenoid support this ; but the birds become smaller southwards instead of larger, and the female shows no sign of the pinkish underparts and barred throat characteristic of the south-western form. It seems more probable that the changes are clinal (p. 187). In the Louisiades H47, the female shows the bright yellow underparts, white throat (though without mottling) and narrow gorget of group E, but has a very brownish mantle like those of group H, while the male has olive edges to the wing quills. This form could equally well be assigned to either of these groups. The female on Tabar H50 resembles those of group E in the same respects, as to a lesser degree does that on Manus H5r1. It seems reasonable to suppose that the inter- gradation in the Louisiades is secondary, and this might be true of the Tabar race also; but it is most unlikely that the small and recently-arrived populations of dahli E25 in the Bismarcks could sufficiently swamp the presumably large population on isolated Manus. This is surely a case of convergence, as in Tonga H57. While the male on Damar H4r is very like those of Timor and Babar H39-40 ee : THE PACHYCEPHALA PECTORALIS SUPERSPECIES 165 (which I suppose to have been affected by gene-flow from group A), the female is different, closely resembling that of the Sula Isles H45 (which has affinities with group B). There may perhaps have been gene-flow across the Banda Sea. But the barred throat characterizing these females is also found in Australia, and the male on Damar has grey edges to the wing quills like those of group E. If gene-flow (other than from group A) is involved here, it seems more likely that it has been across the Timor Sea from northern Australia E23. The three forms in the Santa Cruz archipelago are remarkably different. The male on Ndeni and the islets H52 is melanic and has the throat feathers elongated, while those on Utupua H53 and Vanikoro G38 are close to the standard. Groups E to H differ mainly in the characters of the female. The Ndeni female is very near the standard, and a typical member of group H; that on Vanikoro is quite different, most like the Snow Mountain female E27, and apparently a northern representative of group G; while that on Utupua is intermediate between them. It is possible that this intergradation is primary, and that group H has spread west and east after its origin in the Santa Cruz, but more probably the contact was secondary. Marks of hybridity Knowing that hybridization is possible between very dissimilar forms of P. pectoralis, we have considered many forms which are intermediate, both geographically and phenotypically, between well-marked groups to be of hybrid origin, even though their variability is not exceptional. Several of these presumably hybrid forms differ from both putative parents (and from most other forms in the superspecies) by characters which are retarded in the male, or advanced in the female plumage. Partly olive (or grey) tails in the male are found in P. soror and the southern members of groups E, F and G; in certain presumptive hybrids (San Cristobal C17, Timor to Damar H39-41, Louisiades H47 and Kandavu H54); and nowhere else. Female throats without melanic washes or mottling are found in P. sovoy and the southern members of groups E and G; in some hybrids (Flores Sea and Salayer A4—5, San Cristobal C17, Timor and Babar H39-40, Louisiades H47 and Utupua H53) ; and nowhere else. Partly black tails in the female are found in P. soroy and members of group E ; on San Cristobal C17 (and black shafts on Babar H4o); and nowhere else. The presumptive hybrid females on Obi B8, Babar H4o and Utupua H53 have much brighter yellow bellies than their relatives on either side. In other parts of the species range, the amount of black in the tails of both sexes, and the depth of carotenoid coloration, are involved in character-progressions which may be climatically determined (p. 186). But while in these clines intensity of pigmentation increases with increasing temperature and humidity, in the forms under discussion the male tail tends to be pale and the female tail and belly to be deeply-coloured. Furthermore, there is no reason to suppose that the islands in the Flores Sea, Timor, Babar, Damar, Obi, the Louisiades, San Cristobal, Utupua and Kandavu share environmental factors which distinguish them from neighbouring islands. If the apparent regularities arereal, it seems most probable that they are related to the secondary intergradation which characterizes P. pectoralis in these areas. These trends are not the phenotypic expression of heterosis, since the variable 166 THE PACHYCEPHALA PECTORALIS SUPERSPECIES hybrid populations E26, D18b & 1gb-c do not show them. Possibly the tendency towards breakdown of the sexual dimorphism reflects genetic disturbances produced by hybridization, or selective changes involved in regaining a balanced genome. These might be expected sometimes to blur the distinction between the adult female and juvenile plumages, producing retarded rather than advanced characters in the female. This may be the explanation for the reversal of south-to-north changes in northern Queensland (p. 156). Many characters might be considered as advanced or retarded, and it would be dangerous to take their occurrence as evidence of hybrid origin ; but the possibility of such effects needs to be considered in assessing the systematic importance of characters, and in tracing environmental correlations. Though these characters tend to appear together in forms of hybrid origin, the distribution of each is very erratic. The abrupt loss of black from the male tail (coinciding with a sharp increase in size, and the loss of ventral rufous) at the Ombai Strait, and its progressive reappearance eastwards here (H39 to 41) and in the Louisiades (H47a to c) suggests that this character may be dependent upon a rather precise balance between genes from each parental group. If this is true of all the “ marks of hybridity ’’, and each has a different threshold value which depends also upon the parental genomes involved, the sporadic realization of these trends is not surprising. Sympatry All the forms of the pectovalis superspecies replace one another geographically, except in New Guinea. There P. sovoy and P. schlegelii represent one another altitudinally, but meet at about five thousand feet. There is no sign of intergradation between these very distinct species. They are the endemic representatives of P. pectoralis, and occupy the primary hill and mountain forest respectively. P. pectoralis itself has only been able to enter New Guinea by way of certain disturbed habitats. In the lowlands of the south and south-east E24-25a, it occupies coastal second growth, and may never come in contact with P. soror of the hill forest (though they are only three miles apart on Fergusson and Goodenough Islands respectively). Another race of P. pectoralis is known only from two river valleys on the northern slopes of the Snow Mountains E27. These valleys have been much affected by intensive native cultivation, which has stripped the forest from their floors and far up their sides. The local race of P. pectoralis seems to live largely in second-growth stands of Casuarina in the resultant grasslands, but also in the forest (Archbold, Rand & Brass, 1942). P. schlegelii descends a little way into the valley forest, and is found there side by side with P. pectoralis, with no signs of intergradation. Else- where in New Guinea P. soror is found up to the level of the valley floors, but it seems to be absent locally, and only appears in the hill forest more than a thousand feet lower. P. soroy and the mountain race of P. pectoralis are so much more alike than intergrading forms outside New Guinea that one would expect that they must be conspecific. But the absence of interbreeding between them can scarcely be due solely to extrinsic barriers. They have been collected within a few miles of one another (on the only expedition which has yet encountered the montane race of P. pectoralis) i ee ae a, Se a ee THE PACHYCEPHALA PECTORALIS SUPERSPECIES 167 and must surely meet on occasion. P. Pectoralis in the Snow Mountains is separated from its relatives in southern New Guinea by over three hundred miles of primary forest. It seems possible that further investigation will show the known populations to be a few of many, small and scattered among the mountains, wherever disturbance of the forest by man, landslide, wind, flood or fire provides a shifting foothold (cf. Rand, 1941, on the origin of the grassland avifauna). In any case, P. pectoralis has presumably passed through the range of P. soror to reach the known localities in the Snow Mountains. It is not strictly true to say (Rand, 1940) that this race of P. pectoralis shows no closer relationship to P. soror than does the southern New Guinea one. Discounting the effects in southern New Guinea E24 of gene-flow from group F, most of the changes (male throat feathers grey-based, collar reduced, mantle greener, edges of wing-quills olive and broader, female with olivaceous breast, brown cap and blacker tail) are in fact in the direction of P.soror. But even if these differences SCHLEGELII 3 OOM EET itd iLL ITLT TR Fic. 3.—The three sympatric forms, in the Snow Mountains of New Guinea. P. pectoralis E27 ; P. sovor 2a; P. schlegelii 3a. 168 THE PACHYCEPHALA PECTORALIS SUPERSPECIES from other members of group E imply gene-flow from P. soror, rather than parallel adaptation to higher altitudes (p. 188), the introgression must have been slight and soon ended. The great reservoir of sovor genes would otherwise soon swamp the small populations of P. pectoralis in the mountains, destroying not only the rather slight phenotypic differences but the genetic determination of habitat preference which presumably keeps the two forms apart. Although the evidence is very indirect, it seems that P. soroy and group E of P. pectoralis are genetically sympatric species. A rather similar situation arises in the Bismarcks, where dahli E25 occupies small islands (but also occurs at certain coastal localities on the large islands) while the endemic race H48 keeps strictly to the dense forest inland (Dahl, 1899). Probably because of this difference in habitat preference, there seems to be little interbreeding between the two forms, though Hartert (1926) records that occasional males on New Britain combine the smaller size and greener mantles of the endemic race with the greyer wings of dahli, and Mayr (1955) suggests that much of the internal variation of dahli may be the result of gene-flow from group H. The two forms must meet occasionally, though dahli may not normally breed on the large islands. It would be reasonable to suppose that groups E and H must be partly isolated by intrinsic barriers, like P. soyoy and the Snow Mountains race of P. pectoralis. Yet in the Louisiades H47 the same groups have merged completely. Several explanations might be put forward to account for this anomaly, any or all of which may be true in part. It may be that in the Louisiades existing barriers have been broken down by hybridization (cf. Sibley, 19546), permitting the populations to merge, while dahli has not been long enough in the Bismarcks for this to have happened. Or the populations of group H endemic on the relatively small islands of the Louisiades may have been inadequately shielded from interbreeding by differences in habitat-preference. Or after interbreeding in the Louisiades, dahli in the islands west of the Louisiades (E25b-c) may have secondarily developed barriers against group H, before invading the Bismarcks. In the absence of positive evidence, it is safest to assume that the two stocks in the Bismarcks represent one another geographically, and seldom meet in the breeding season. Yet it is conceivable that intrinsic barriers to interbreeding are effective here. From this, and from the different reactions of the schlegelii and soror assemblages (and of differentiated stocks within the latter) when they meet in New Guinea and in the archipelagos, it is clear that species-limits need not at all closely follow the pattern of descent and resemblance. P. schlegelii, P. soroy and group E of P. pectoralis are genetically sympatric (Cain, 1953) and must be regarded as good species. Yet the subspecies-groups A to H of P. pectoralis, some of them much less alike than are P. sorvor and group E, are inter- connected by a web of secondary intergradation (Text-fig. 4). In the absence of evidence that groups E and H meet in the breeding season without significant interbreeding, all these groups can be included in a single species. GROUP AFFINITIES The schlegelii assemblage P. schlegelt, groups A to D of P. pectoralis, and P. flavifrons all differ markedly from the standard patterns for the superspecies. Each is distinguished from the THE PACHYCEPHALA PECTORALIS SUPERSPECIES 169 others by characters which are not merely striking but stable and systematically important (as is shown by their co-variation). At first sight they seem to be indepen- dently derived from more standard forms, rather than closely related to one another. There is little in common, for example, between P. schlegelii and the Fijian group D (Text-figs. 3 and 2, bottom). Yet there are important resemblances between members of this assemblage of forms. Most striking is the similarity between females of P. schlegelii and of P. SCHLEGELII SOROR A PECTORALIS E B ate F c a G D FLAVIFRONS Fic. 4.—Diagram of the phylogeny suggested for the superspecies, showing secondary intergradation between subspecies-groups of P. pectoralis (dotted) and sympatric co-existence in New Guinea (arrows). 170 THE PACHYCEPHALA PECTORALIS SUPERSPECIES pectoralis in the northern Moluccas (group B). Furthermore, the males of the same forms are alike in having black chins, and throat-patches overlapping the gorgets without black tips to the feathers. These two characters of the males are shared by forms in most of the groups of the schlegelii assemblage. The black chin is absent from all the hybrid forms and from group D, but characterizes P. schlegelii, P. flavi- frons and undiluted members of groups A to C. It is not found elsewhere in the superspecies (except in the black-throated race H57). The unusual structure of the gorget-edge is less affected by gene-flow, and is found in almost all the members of the assemblage which have gorgets. Elsewhere it occurs only on New Caledonia G34 (possibly related to the assemblage) and Ndeni H52 (see p. 162). Several other characters serve to link two or more groups. In this way each of the groups is united with at least one other: groups A to C of P. pectoralis to one another and to P. schlegelii, group D to group C, and P. flavifrons to group D. Table III sets out the main characters which occur in more than one group of the assemblage, while remaining rare elsewhere in the superspecies. The table does not illustrate the resemblance between females of P. schlegelii and of group B, since this depends on the conjunction of several characters (grey-barred throat, olive breast, grey auriculars and lack of phaeomelanins) which occur separately in several different forms outside the assemblage. TasieE III.—Characters Uniting the schlegelii Assemblage. schlegelit assemblage sovor schlegeltt A B Cc D flavifrons assemblage Male: Chin black . 0 7 L3 zr 6-7 9-15 — x 6 a Throat yellow . _ = —_ — 9-17 18-21 x a Jak (E26) No gorget . 5 _ oo —_ 12 18-19 x 5 — gorget edge : 5 TS 8} 1-3 6-8 9-II, — - H52 (H43) 13-15 breast rufous 5 - 3 1-4 — 17 — — . —_— (G34) Forehead yellow . _ —_ _— — 18-109, x 5 = 21 Mantle melanic . _ 4 — 13-14 20 x Sz (H55-56) Wing black : 5 TER 4 — 13-14 — x : — Tail black . 0 - i3 I 6-7 13 — x f H52 Female : Cheeks yellowish . = _ — 9,11 &12 18-19 — 6 —_— Underparts streaked . — _ —_— 9-16 18-19 —_— 5 _— Wing russet : _ — — 9-16 18-21 — _ The soror assemblage P. soror and the remaining groups of P. pectoralis are united negatively rather than positively, by the absence of those unusual characters which distinguish and link ——— ew THE PACHYCEPHALA PECTORALIS SUPERSPECIES 171 together the groups and species belonging to the schlegelii assemblage. Since group H asa whole scarcely departs from the average characters of the whole superspecies, it can be associated with other groups only negatively. However, there are a few characters and trends which link P. soror with groups E to G and these with one another, and are rather rare in the schlegelii assemblage. These are set out in Table IV. TaBLe IV.—Characters U muting the soror Assemblage (except Subspecies-group H). soror assemblage —— a schlegelit sovor E F G assemblage Male : Carotenoid pale 5 : 1-3 22 28-30 34, 36-38 —_— Quills grey 5 5 “ I-3 22-25 28-30 — - Dro, 21 Coverts yellow . 5 0 —_— 22-25 28-32 35 - Dr8-19 Tail pale . 0 6 I-3 22 28-32 B4—S7N te C17 Female : Throat white . 2 . I-3 22-27 — 34-38 Ct A4-5 Throat streaked (not belly) —_— 25,27 — 38 A — Cap grey (not auriculars) . — 22-25 28-33 — 5 A5 Several of these characters seem to appear chiefly in higher latitudes, and may be related to the generally cooler climates encountered by these groups. Yet they are very different from the characters of P. schlegelit, which lives at higher altitudes than P. soror. The resemblances between group F and the Vanua Levuan race Dig ‘(grey-edged wings and yellow-edged wing-coverts in the male; barred throat, melanic belly and almost complete absence of carotenoid in the female) may be due partly to selection under cooler climates. These characters, and others which (it may be objected) could be used to erect any number of alternative assemblages, are evidently unstable, since they appear independently in widely-scattered localities and cut across the discontinuities indicated by the co-variation of more stable characters, ARRANGEMENT The considerations of relationship, intergradation and sympatry discussed above permit the various forms to be arranged systematically. The categories of super- species, species, subspecies-group and subspecies are used, and considerable infra- subspecific geographical variation is recognized. The superspecies The category of Artenkreis or superspecies (Rensch, 1928; Mayr, 1931a@) was introduced for groups of strictly allopatric representatives, some of which are too unlike to be considered conspecific. Mayr pointed out that the great practical usefulness of this category might be expected to decline as the possibility of unlike Tepresentatives being conspecific became generally accepted. More recently the scope 172 THE PACHYCEPHALA PECTORALIS SUPERSPECIES of the superspecies has been extended to include groups of forms some of which are genetically sympatric (Cain, 1953), with their breeding ranges even overlapping slightly (Mayr and Vaurie, 1948 ; Mayr, 1949; and definition in Mayr, Linsley and Usinger, 1953). This extension gives the category permanent value, in expressing the relationship between forms which can never be considered conspecific yet which have barely ceased to represent one another geographically. But it removes the objective criterion which limited the use of the superspecies under the older usage, and opens the way to further extension. There is a danger of the category losing its special connotations of geographical replacement, becoming synonymous with the species-group. Sometimes a species or superspecies is represented locally by a pair of sympatric species, so that it would be arbitrary to include one rather than the other in the superspecies, or to exclude both. Cain (1954) has introduced the term ‘“ doublet ” for such pairs in the Pitilinopus purpuratus superspecies, and this usage is helpful in indicating cases of double invasion. Where triple invasion by a single species has resulted in three representatives locally, the term “‘ triplet’? may be used. For example, Zosterops lateralis is represented by a doublet on Lord Howe (Z. strenua and Z. 1. tephropleura), and by a triplet on Norfolk Island (Z. albogularis, Z. tenuirostris and Z. /. norfolciensis). P. soror is so like many members of P. pectoralis, while P. schlegelii is so strikingly different, that the former is the obvious New Guinea representative. P. soror and P. pectoralis seem to be only genetically sympatric, without actual overlap, and might be included in one superspecies. But we have seen that, while P. soror is indeed the more closely related to some subspecies-groups of P. pectoralis, P. schlegelii is so to others. The two species should therefore be considered as a New Guinea doublet, like Ptilinopus coronulatus and Pt. pulchellus (Cain, 19546). The local intrusion of P. pectoralis itself means that there is a triplet on the northern slopes of the Snow Mountains. Species All the remaining forms are geographical representatives. Those that intergrade must be conspecific, so that even the extreme forms on Sumba and in the northern Moluccas, Solomons and northern Fiji belong to P. pectoralis. Certain striking single characters have evidently arisen several times independently, and most modern systematists would agree in placing within P. pectoralis both the hen-feathered races on Rennell and Norfolk Island C16 & F33, and the melanic ones in the Solomons and Santa Cruz C13 & H52. However, there are a few extreme forms (Text-fig. 5) whose status is doubtful, and can only be decided by comparing the degree of difference between them and their nearest relatives with that shown between sympatric species on the one hand, and between intergrading subspecies on the other. But in the pectoralis superspecies, two sympatric forms are much more alike than many intergrading ones: the relationship between visible differentiation and the establishment of barriers to interbreeding is not the same in New Guinea as in the archipelagos. The forms of doubtful status are all found on more or less isolated islands, whose avifaunas are Iya) 2 hE, » THE PACHYCEPHALA PECTORALIS SUPERSPECIES 173. much poorer in species than those of New Guinea and include at most one other species of Pachycephala (P. rufiventris on New Caledonia). It is reasonable to suppose that the intergradation of unlike subspecies in the archipelagos is more relevant here than the coexistence of like species in New Guinea (p. 179). P. flavifrons in Samoa is a derivative of the Fijian group D (Table III), but deserves the specific rank always accorded it. Group D is itself the most aberrant in P. pectoralis, and is probably on the borderline of ethological and genetic incompatibility with the rest of the species, while P. flavifrons has acquired further unusual characters —including the unique cock-feathering of its females. Previous authors have considered the Tongan form H57 as a distinct species (P. melanops) because of the black throat of the male. In other ways this form is FLAVIFRONS is — AE ty GUE age Fic. 5. Three unusual island forms. Samoa, P. flavifrons; Tonga, P. p. melanops H57; New Caledonia P. [p.] caledonica G34. 174 THE PACHYCEPHALA PECTORALIS SUPERSPECIES rather close to the standard pattern. It seems to be a member of group H, not more modified than might be expected from its isolation and the poverty of the avifauna in which it is placed. There is no reason to believe that the black throat reflects any very profound genetic change. The white throat-patch is very restricted on Sumba At and in P. schlegelii, and reduced to narrow white barring on black in individuals of the latter’s close relative P. auvea. The mountain species P. nudigula and P. implicata (derived from the pectoralis superspecies) have acquired blackish throats independently. On the other hand, the black throat does alter the appearance of the male most markedly, and might therefore be of ethological importance. But so does the abolition of the gorget, and even more the total loss of the distinctive male pattern: yet there has been gene-flow between races with and without gorgets, while hen-feathered males in dimorphic races are able to find mates and breed. It is most improbable that the black throat alone would be sufficient to restrict inter- breeding while further barriers were selected for, in the event of a second invasion of Tonga. ‘‘ P. melanops’’ ought therefore to be considered as a race of