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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
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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
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‘ 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
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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 ”’.
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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
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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).
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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
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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.
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THE FAMILY TRICHIURIDAE
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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
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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
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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<