A MtieucanJMllsellm PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N. Y. I0024 NUMBER 24 I 0 MARCH 25, I970 The Hyobranchial Apparatus of Teleostean Fishes of the Families Engraulidae and Chirocentridae BY GARETH J. NELSON1 In a previous study of the gill arches of teleostean fishes of the family Clupeidae, the writer defined certain phyletic trends, and on this basis erected an hypothesis of the phyletic interrelationships of the currently recognized subfamilies (Nelson, 1967a, fig. 9). To check these results, study of the gill arches is extended, in the present paper, to the families Engraulidae and Chirocentridae, which probably include the nearest Recent relatives of the Clupeidae. In addition, the structure of the branchiostegal apparatus is reviewed for all three families. On the basis of the new data, it is possible to arrive at a more precise conception of the phyletic interrelationships of the main groups of clupeiform fishes, and to propose a revised higher classification: ORDER CLUPEIFORMES SUBORDER DENTICIPITOIDEI SUBORDER CLUPEOIDEI SUPERFAMILY CHIROCENTROIDAE SUPERFAMILY ENGRAULOIDAE SUPERFAMILY PRISTIGASTEROIDAE SUPERFAMILY CLUPEOIDAE 1 Assistant Curator, Department of Ichthyology, the American Museum of Natural History. 2 AMERICAN MUSEUM NOVITATES NO. 2410 MATERIAL AND METHODS Specimens available included representatives of virtually all genera commonly recognized (e.g., by Norman, 1957; Hildebrand, 1943, 1963a, 1963b; Svetovidov, 1963; Whitehead, 1968a). They were generally small- to-medium adults. The hyobranchial apparatus was dissected as a unit from a given specimen, stained with alizarin (in aqueous 2% potassium hydroxide), and cleaned and examined under a binocular microscope. Drawings were prepared either freehand, with the aid of an ocular micrometer, or by tracing from photographs. Terminology follows that of Nelson (1969). Gill-arch dentition was noted for at least one young adult per species (table 1). Teeth were recorded as being present when one or more could be observed. Branchiostegal rays were counted on both sides of the same specimens, with the highest number recorded (table 2). Some BHP BH B1 B2 B3 B4 BC 82 -122 83 H3 FIG1.Enrui3odx,glrhs .Bairnha eis aea iwo 84 Elw5Cl-5 El-4 B C 0 FIG. 1. Engraulis mordax, gill arches. A. Basibranchial series, lateral view of left side. B. Ventral parts, ventral view. C. Dorsal parts, ventral view. D. Dor- sal parts, dorsal view. Cartilage stippled, bone clear. Cartilaginous articular areas at tips of ceratobranchials and epibranchials not shown. Basihyal omitted in B. E4-5 tilted backwards in D. Abbreviations: B1-4, basibranchials 1-4; BH, basihyal; BHP, tooth plate over- lying basihyal; C1-5, ceratobranchials 1-5; E1-5, epibranchials 1-5; H1-3, hypobranchials 1-3; I1-4, infrapharyngobranchials 1-4; UP4-5, upper pharyn- geal tooth plates 4-5. 0 - c~cl u 0l - CZ 3 14% C ,**% u--' -4 U-0 ,3C LO C) LL ) -0N Lr 0 c u qh4w_dI %e. - CY b*szoAd !,l _ -%s V C.^) . i aeop ~~s v ~Csm CN Ce) q% %& 14% ON-1. 4 -400 LA01 sa C,,O LLJ\V .0F LLJ -,N ,t .t U * o '- la LLi -4wwwlpp~_ m LLJ q% Ih,- LLJ V .~ C - Cr) 4 U) q%b -qm C-t LU 00, LC= 400, Ago 04 CY) q,, -2 U-) (-) 4 q"%b q%46 qq% .id 11~0 amw/", C3* "Wp_ _ _ 40 -..ddo LLJ N - C14 m 14 U-) < "I"% *4%%*4 6 AMERICAN MUSEUM NOVITATES NO. 2410 specimens had different counts on each side, the left side usually having one ray more than the right. The counts are given as posterohyal ("epihyal") rays + anterohyal ("ceratohyal") rays. For cases in which the anterohyal had a distinct posteroventral process supporting a single ray, the count is given as posterohyal rays + this single ray + the remaining anterohyal rays. The writer is indebted to the following persons and institutions for making specimens available: Drs. W. D. Anderson, Jr. and C. R. Gilbert, University of Florida; Mr. G. S. Arita and Dr. C. C. Lindsey, University of British Columbia; Drs. J. E. Bohlke and J. C. Tyler, Academy of Natural Sciences of Philadelphia; Dr. E. J. Crossman, Royal Ontario Museum; Dr. W. C. Freihofer, Stanford University; Dr. P. H. Green- wood and Mr. P. J. P. Whitehead, British Museum (Natural History); Dr. G. W. Mead, Museum of Comparative Zoology; Mr. L. P. Woods, Field Museum of Natural History. Part of the work was done at the Swedish Museum of Natural History and part at the British Museum (Natural History) during 1966-1967 with support from a National Science Foundation Postdoctoral Fellowship (No. 46007). RESULTS ENGRAULIDAE: Gill-arch structure is relatively uniform throughout the family, therefore only the gill arches of Engraulis are figured (fig. 1). Like those of clupeids (Nelson, 1967a, p. 394) the dorsal parts of the arches tend to be in contact along the dorsal midline, except in the examined species of Coilia (fig. 3A; see also Joshi and Bal, 1953, fig. 8). Whether or not in contact, the second and especially the third infrapha- ryngobranchials of each side have an elongate, anteriorly directed medial process, a feature by which gill arches of clupeoid fishes may be recognized at a glance. Unlike clupeids, the engraulids have the basihyal cartilaginous and rudimentary; the second basibranchial is the longest endoskeletal ossification of the basibranchial series; and dermal tooth plates generally occur in abundance over most of the gill-arch surfaces (figs. 2-4). The tooth plates include a basihyal tooth plate (BHP), which if present at all is severely reduced, and tooth plates fused with basibranchials 1-3 (in some species, however, one or more basibranchials are secondarily toothless). Tooth plates of irregular size and shape some- times occur over the fourth basibranchial. "Lower pharyngeals" are almost always present, composed of tooth plates fused with the fifth ceratobranchials. Dorsally, the dermal skeleton is well represented by paired series of tooth plates, some free and others fused with their endoskeletal supports. Five pairs usually occur (figs. 3-4; Nelson, 1969, 1970 NELSON: TELEOSTEAN FISHES 7 Bi BHP B4P -Hi B2WB2 ~~~BH Bi B2 B3 BI. -H2 A B41''7 \H3 i-I B4 ~~~~~12 12 13 ~~~~~~~13 UP4-5~ \J I'1 \\~1 B C1-5 C El-4 D El-4 FIG. 5. Chirocentrus dorab, gill arches. A. Basibranchial series, lateral view of left side. B. Ventral parts, ventral view. C. Dorsal parts, ventral view. D. Dor- sal parts, dorsal view. Abbreviations: Bl-4, basibranchials 1-4; B4P, tooth plate overlying basibran- chial 4; BH, basihyal; BHP, tooth plate overlying basihyal; C 1-5, cerato- branchials 1-5; El-4, epibranchials 1-4; Hl-3, hypobranchials 1-3; I1-4, infra- pharyngobranchials 1-4; UP4-5, upper pharyngeal tooth plates 4-5. pl. 82, figs. 4, 5). The anterior pair is supported by the anterior ends of the first epibranchials. In most Indo-Pacific genera these tooth plates are free, but in Engraulis, some species of Stolephorus, and in virtually all examined American engraulids, they are fused with the epibranchials, a feature which so far as is known does not characterize any other group of Recent teleostean fishes. Posteriorly, tooth plates are almost always fused with the second and third infrapharyngobranchials. In- variably present, more posteriorly, are paired tooth plates, one pair (UP4) underlying the cartilaginous fourth infrapharyngobranchials, and one pair (UP5) partially underlying the fourth epibranchials. CHIROCENTRIDAE: Gill arches of Chirocentrus (fig. 5) have the dorsal parts well separated in the midline. The second and especially the third infrapharyngobranchials have long medial processes. The basihyal is small, but ossified and tilted downwards. Dermal tooth plates are well developed: one (BHP) over the basihyal, one fused with the second basibranchial, one (B4P) over the cartilaginous fourth basibranchial, and paired plates fused with the fifth ceratobranchial; dorsally, how- LO H- HH- - HH- - -4F4E4E-HH H - E-HHHHHH4-I~- dz H HH HbHHHHHHHHHHHHH co PC4 -<1 HHIo Hl H 0 C-,~~~~~~C C~~~~~~~~~~~C 0 0~~~~ G d , w|<> i 0 neo Wt 8 H-E E4FIEHHHHHHH4E--F- E-HHHHHHHHHHHH--4P- ~4E- E0 - HE- H HH H H H HH H H HHH H HHH H H C\i~~~ ~ ~ Q i Q @ Q f +>vv ~~~~~~~IIII II II I H l~~~~~~~~~~~~~t I m 9 LrI [- HHHHoo HHHHHHF HH F-4E-HHoH- c)o HH HE H HHoo X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X lX lX lX lX X X X X X X X X _; .3 X X X X X X X X X E- I X - X - X HH~-o . FE4 E-4 E- EH H 'ZI '..) -ts Zs Q . .- biD v~~~~~~0 4~~~~ 10 LOFr) FH H HHE4E49H H H -4E-H E-4HHE-4 E--HE-4 .-H F-H E-4 .-H F-H E-1 HF--4 E-4 04 P4 H o0 o o o 0 0-C ) x x x I I x x x x X lII 11 I I &-, x x x x x x x x I x x x x x x EH o Ix I I FHFH0o HH-4EHH-o Ho HHoo -oHHHH-H wzp t!~~~~~~~~U o t3ts -,Z~~~ C 11 H o o000 o 0000o 0 0 00000000oo UQ u I I to *ld -Z. I I I Ms U CO U = I I I I *_tU O X c>o _O X1I11I11I1 II -o 0 0 > I I I I I I I I C3 I I I I 0 I I I I I -o o >z Xl = H H H o o o o 0000000000oo Q CZ 4 - 0 0 0~ U "O' 0 WO Q, - S 0~~~~~~~~~ 0~~~~~~~ o ~ ~ ~ ~ 4-1 Lr o t 13 12 1970 NELSON: TELEOSTEAN FISHES 13 ever, the only dermal elements represented are the upper pharyngeal plates (UP4-5).