-2 V C. ..>.«./-. i. 19 rH , LIBRARY (NATURAL HISTORY) ZOOLOGY Vol. 20 1970-1971 BRITISH MUSEUM (NATURAL HISTORY) LONDON: 1975 DATES OF PUBLICATION OF THE PARTS No. I . . ... 18 August 1970 No. 2 . . . . •; . 9 November 1970 No. 3 ..... 20 January 1971 No. 4 . . . . .10 December 1970 No. 5 . . . .31 December 1970 No. 6 . . . . .21 December 1970 No. 7 . . . . .28 December 1970 No. 8 . . . . . 8 February 1971 Printed in Great Britain by John Wright and Sons Ltd. at The Stonebridge Press, Bristol BS4 5NU CONTENTS ZOOLOGY VOLUME 20 PAGE No. i. The clupeoid fishes described by Steindachner. By P. J. P. WHITEHEAD (Pis. 1-3) ........ No. 2. The type specimens of Sipuncula and Echiura described by J. E. Gray and W. Baird in the collections of the British Museum (Natural History). By MARY E. RICE and A. C. STEPHEN (Pis. 1-3) No. 3. The types and figured specimens of Unionacea (Mollusca : Bivalvia) in the British Museum (Natural History). By R. I. JOHNSON (Pis. 1-2) No. 4. A review of the species of Hemilepistus s. str. Budde-Lund, 1885 (Isopoda, Porcellionidae). By R. J. LINCOLN .... No. 5. A taxonomic revision of the oligochaete genus Eukerria Michaelsen, 1935 (Ocnerodrilinae, Megascolecidae) . By B. G. M. JAMIESON No. 6. Observations on the electra dolphin, Peponocephala electra. By W. H. DAWBIN, B. A. NOBLE and F. C. ERASER .... No. 7. The species of Macrophthalmus (Crustacea : Brachyura) in the collections of the British Museum (Natural History). By R. S. K. BARNES ........... No. 8. Observations on the systematics of nematodes belonging to the genus Syphacia Seurat, 1916. By C. G. OGDEN (Pis. 1-5) . Index to Volume 20 . 47 73 109 173 203 253 281 THE CLUPEOID FISHES DESCRIBED BY STEINDACHNER P. J. P. WHITEHEAD BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 20 No. i LONDON : 1970 THE CLUPEOID FISHES DESCRIBED BY STEINDACHNER BY PETER JAMES PALMER WHITEHEAD Pp. 1-46 ; 3 Plates, 4 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 20 No. i LONDON: 1970 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, is issued in five series corresponding to the Departments of the Museum, and an Historical series. Parts will appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department. This paper is Vol. 20 No. i of the Zoological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation Bull. Br. Mus. not. Hist. (Zool.). Trustees of the British Museum (Natural History), 1970 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 18 August, 1970 Price £i qs. THE CLUPEOID FISHES DESCRIBED BY STEINDACHNER By P. J. P. WHITEHEAD INTRODUCTION MOST of the new fishes described by Franz Steindachner (1834-1919) are in the Naturhistorisches Museum in Vienna ; a few types and many duplicates are found in other museums. Steindachner's original descriptions and figures are usually excel- lent, often with far more detail than his contemporaries troubled to include, but modern studies have made a reassessment of his species urgent in certain groups. The opportunity is taken here to redescribe and discuss the types of Steindachner's 21 clupeoid fishes (i Dussumieriidae, 12 Clupeidae, 8 Engraulidae — see Table i). Between 1859 an(^ I9I7» Steindachner produced nearly two hundred and fifty papers in which he described about a thousand new species of Recent fishes, 29 fossil species and a number of new reptiles and amphibians. Most of the specimens re- sulted from his participation in several expeditions, the most important in the present study being those to South America and West Africa. As a student, Steindachner's legal studies had given way to an interest in natural history (and ichthyology in particular), largely through the encouragement of Eduard Suess. By 1857 Steindachner was a regular visitor to the Imperial natural history cabinet and in 1861 he accepted a permanent post there. One of his first tasks, suggested by Rudolf Kner, was to share with Giovanni Canestrini the study of the material brought back from the Novara Expedition of 1857-9. Determined to enlarge the Imperial collections, Steindachner made expeditions to Switzerland, southern Spain, Portugal and the Canary Islands and in 1868-9 he collected in Senegambia. Subsequently, he accepted Louis Agassiz's invitation to come to Cambridge (Massachusetts) to work on the Thayer material collected in Brazil in 1865-6. Granted leave of absence from Vienna, Steindachner joined Agassiz in March, 1870 and later agreed to accompany him on a collecting trip in the ship Hassler on a cruise down the Atlantic coast of the Americas and back up the Pacific coast to San Francisco, a voyage of nine months. For a further seven months Steindachner collected in the United States for the Vienna museum, before returning home. Twenty-nine years were to elapse before he revisited South America, this time to the northeast provinces of Brazil. In the meantime he participated in and later led three of the Austrian deep-sea Mediterranean expeditions and two Red Sea expedi- tions, and the fish collections at Vienna grew at such an enormous rate that new accommodation was required. The transfer of the fish and reptile collections from the seven small and dark rooms of the Imperial Cabinet in Joseph Platz to the present museum building was completed by 1886 and the following year Steindachner was 4 P. J. P. WHITEHEAD appointed Director of the zoological collections with a suite of rooms which now houses the ichthyological collections. A full account of Steindachner's career has been given by Kahsbauer (1959). Perusal of the modern fish collection shows to what extent it is indebted to Stein- dachner's efforts. Amongst the material, there are also Brazilian specimens collected by Johann Natterer (recognizable by red painted roman and arabic numerals on the backs of mounted skins), Japanese fishes (dried) from Burger (many still with paper glued to protect the fins), Red Sea fishes from Eduard Riippell, type and other material relating to Rudolf Kner and Johann Heckel, and exchange specimens from Leyden, Paris, etc. Kahsbauer (1959) gave a useful index of the new species described by Steindachner (a few species missing and some MS. names or already published names included). The Steindachner types are not segregated from the non-typical material and are not always indicated as such ; in some cases a type indication relates merely to a MS. name. No peculiarities of labelling, type of bottle, marking of specimens, etc., were found which would aid in the recognition of types other than the dates given on the labels (which may in some cases refer to the date of incorporation and not to the date of collection). For the 21 clupeoid species, types were not found for the following species : Clupea rechingeri ( = ? Herklotsichthys punctatus) Clupea notacanthoides ( = Ethmidium maculatum notacanthoides) Engraulis natter eri ( = Anchoviella natter eri} Engraulis poeyi ( = Lycengraulis poeyi) The types of 2 species were reported by Steindachner (see under species) to be in Stuttgart : Clupea macrolepis (SMNS. 2292) Clupea neopilchardus (not found, see p. 16). As an indication of Steindachner's general approach to ichthyology, his clupeoid species can be said to have been described accurately, usually at greater lengths than in contemporary descriptions, but with little or no attempt to explore affinities beyond those of immediate specific relationships. Initially, his clupeoid genera followed those of Valenciennes (1847, 1848), but he later tended to use the two com- pendium genera, Clupea and Engraulis, favoured by Giinther (1868). Fourteen Steindachner names are here recognized as senior synonyms (Table i). The following abbreviations have been used : S.L. standard length tot.l. total length g.r. gillrakers NMV Naturhistorisches Museum, Vienna SMNS Staatliches Museum fur Naturkunde, Stuttgart BMHN British Museum (Natural History), London MNHN Museum National d'Histoire Naturelle, Paris RMNH Rijksmuseum van Natuurlijke Historic, Leyden ZMA Zoologisch Museum, Amsterdam ZMC Zoologisk Museum, Copenhagen STEINDACHNER'S CLUPEOID FISHES 5 I am indebted to Dr. Paul Kahsbauer for his kindly help during my visit to Vienna and for allowing me to borrow many types for further examination. TABLE i Clupeoid species described by Steindachner Steindachner name DUSSUMIERIIDAE 1. Alausa alburnus Kner & Steind., 1866 CLUPEIDAE 2. Clupea rechingeri Steind., 1908 3. Clupea brasiliensis Steind., 1879 4. Clupea macrolepis Steind., 1879 5. Clupea amazonica Steind., 1879 6. Pellonula bahiensis Steind., 1879 7. A lausa fimbriata Kner & Steind., 1879 8. Clupea neopilchardus Steind., 1879 9. Clupea setosa Steind., 1869 10. Clupea notacanthoides Steind., 1869 11. Pellona furthii Steind., 1875 12. Pellona panamensis Steind., 1875 [Pellona staudingeri, MS. name] [Pellona macrolepis, MS. name] 13. Pristigaster (Odontognathus) panamensis Steind., 1876 ENGRAULIDAE 14. Engraulis vaillanti Steind., 1908 15. Engraulis natter eri Steind., 1879 16. Engraulis januaria Steind., 1879 17. Engraulis nasus Kner & Steind., 1866 18. Engraulis peruanus Steind., 1879 19. Engraulis panamensis Steind., 1875 20. Engraulis macropolepidotus Kner & Steind., 1865 21. Engraulis poeyi Kner & Steind., 1865 Identification Spratelloides delicatulus (Bennett, 1831) ? Herklotsichthys punctatus (Riippell, 183?) Sardinella brasiliensis (Steind., 1879) Escualosa thoracata (Val., 1847) Rhinosardinia amazonica (Steind., 1879) Rhinosardinia bahiensis (Steind., 1879) Sardinops sagax sagax (Jenyns, 1842) Sardinops sagax neopilchardus (Steind., 1879) Ethmalosa fimbriata (Bowdich, 1825) Ethmidium maculatus notacanthoides (Steind., 1869) I lisha furthii (Steind., 1875) I lisha furthii (Steind., 1875) Pellona flavipinnis (Val., 1837) Pellona flavipinnis (Val., 1837) Odontognathus panamensis (Steind., 1876) Anchoviella vaillanti (Steind., 1908) Anchoviella nattereri (Steind., 1879) Anchoa januaria (Steind., 1879) Anchoa nasus (Kner & Steind., 1866) Anchoa nasus (Kner & Steind., 1866) Anchoa panamensis (Steind., 1875) Anchovia macrolepidota (Kner & Steind., 1865) Lycengraulis poeyi (Kner & Steind., 1865) Family DUSSUMIERIIDAE SPRATELLOIDES Bleeker, 1851 Spratelloides Bleeker, 1851, Natuurk. Tijdschr. Ned. Ind., 2 : 214 (Type : Clupea argyrotaeniata Bleeker = Clupea gracilis Temminck & Schlegel) . In a recent revision of the genus (Whitehead, 1962) two species were recognized, 5. gracilis (Temm. & Schl.) and S. delicatulus (Bennett). Subsequent studies have dealt with type specimens (Whitehead et alii, 1966 : 33-37), further differences between the two species (Whitehead, 1965 : figs. 2, 3 ; supra-maxillary shape, pos- terior frontal fontanelles) and distribution (Whitehead, 6 P. J. P. WHITEHEAD i. Alausa alburnus Kner & Steind., 1866 = Spratelloides delicatulus (Bennett, 1831) (Plate la) Clupea delicatula Bennett, 1831, Proc. zool. Soc. London, 1 : 168 (Mauritius). Alausa alburnus Kner & Steindachner, 1866, Sitzb. K. Akad. Wiss. Wien, 54 : 387, pi. i, fig. 16 (" Valparaiso " — in fact Samoa, see below). LOCALITY. As pointed out by Giinther (1909 : 384), the original reference number and the locality recorded for this species by Schmeltz (1869 : 25) in the catalogue of the Godeffroy Museum at Hamburg were attributed by Kner & Steindachner to Alausa fimbriata, and vice versa. The correct locality for Alausa alburnus is thus Samoa ; Stolephorus delicatulus is not recorded from the Pacific coasts of America. TYPE MATERIAL, a. LECTOTYPE, a fish of 43-0 mm S.L. ex Godeffroy Museum Reg. No. 2152, from Samoa in 1866, NMV.4282. b. PARALECTOTYPE, a fish of 40-8 mm S.L. (source as above). There are two further specimens, 49-I.-5I-5 mm S.L. ex Godeffroy Museum, from Samoa in 1869, NMV.4328 (locality correctly stated as Samoa on label). DESCRIPTION. A fish, 43-0 mm S.L., LECTOTYPE, ex Samoa ; in fair condition but dorsal and anal fins slightly damaged and caudal lobes broken, NMV-4283. Br.St. ?, D ii 8, P i u, V i 7, A ii 8, g.r. 30. In percentages of standard length : body depth 18-8, body width 9-1, head length 25-6 ; snout length 7-0, eye diameter 7-4, length of upper jaw 9-3, length of lower jaw 107 ; pectoral fin length 12-6, pelvic fin length 10-9, length of anal base 8-1 ; pre- dorsal distance 47-5, pre-pelvic distance 56-5, pre-anal distance 79-5. Body a little compressed, its width twice in depth, belly rounded and without scutes except for W-shaped pre-pelvic scute ; head longer than body depth ; eye diameter a little greater than snout length. Upper jaw reaching to anterior eye border ; pre-maxillae triangular, toothless ; maxillae toothless ; two supra-maxillae, the ist (anterior) plate-like and attached to upper edge of maxilla, the 2nd (posterior) supra-maxilla with slender anterior shaft and expanded posterior part, the latter as deep as long (about i-o mm), its upper profile rising steeply, its lower profile joining anterior shaft opposite that of upper profile (as in Sardinella ; cf . the more asymmet- rical shape in Harengula). Lower jaw about twice as long as deep, highest point in first third of length ; articulation of lower jaw below vertical from anterior pupil border. Posterior border of operculum with slight indentation in upper part ; lower border of operculum horizontal. Cleithrum with well developed fleshy cleithral lobes. Isthmus silvery, sterno-hyoideus muscle ending abruptly anteriorly, the urohyal exposed in front of this until concealed by gill membrane. Gillrakers long, slender, about \ eye diameter, lined with 20-25 fine serrae on each side ; about 6 short gill- rakers on posterior face of 3rd epibranchial ; gill filaments about $ length of gillrakers. Pseudobranch present, exposed, about $ eye diameter. Dorsal surface of head without striae, posterior frontal fontanelles together almost circular, 0-9 mm long (equal to pupil), anterior extension of supra-occipital slender. STEINDACHNER'S CLUPEOID FISHES 7 Dorsal fin origin nearer to snout tip than to caudal base by just over i eye diameter. Pectoral fin tips fail to reach pelvic base by 2^ eye diameters. Pelvic fin base under 8th branched dorsal ray, nearer to anal origin than to pectoral base by f eye diameter. Anal fin origin nearer to caudal base than to pelvic base by £ eye diameter. Unexposed portion of scales with one main and 2 (anterior) to 4 (posterior) sub- sidiary complete vertical striae ; exposed portion of scale without crenulations, striae or perforations. Colour : upper £ brown, remainder silvery ; no silvery lateral stripe. IDENTIFICATION. The absence of a silver lateral stripe (Whitehead, 1962 : 338) and the shape of the posterior frontal fontanelles and posterior supra-maxilla (Whitehead, 1965 : figs. 2, 3), clearly identify the present specimen as Spratelloides delicatulus. The smaller pectoral fins place it in the nominate subspecies (White- head, 1962 : 347), which is consistent with its presumed provenance (Samoa). Fowler (1941 : 565) recognized Alausa alburnus as a distinct species, but Bertin (1943) and Schultz & Wellander (1953) correctly placed it in the synonymy of S. delicatulus. Family CLUPEIDAE HERKLOTSICHTHYS Whitley, 1951 Herklotsichthys Whitley, 1951, Proc. Roy. zool. Soc. N.S.W., 1949-50 : 67 (Type : Harengula dispilonotus Bleeker). A single Steindachner species, Clupea rechingeri, is rather doubtfully included in this Indo-Pacific genus. 2. Clupea rechingeri Steindachner, 1908 = ? Herklotsichthys punctatus (Ruppell, 1837) Clupea punctata Riippell, 1837, Neue Wirbelth., Fische : 78, pi. 21 (2) (Red Sea). Clupea rechingeri Steindachner, 1908, Sitzb. K. Akad. Wiss. Wien, 115 (i) : 1424 (Upolu, Samoa). TYPE MATERIAL. The two specimens described by Steindachner (no size given) cannot now be found. IDENTIFICATION. The criteria now used to distinguish the genera Sardinella and Herklotsichthys (Whitehead, ig64a) do not appear in the description of C. rechingeri and Steindachner gave no clue to its relationship to other species. Regan (1917 : 392) and Fowler (1941 : 597) identified Clupea rechingeri with Harengula vittata ( = Sardinella melanura (Cuvier) fide Whitehead, 19670. : 66). The latter has very distinctive black caudal tips (retained in alcohol specimens), whereas Steindachner's description seems to allude to the general darkening of the whole caudal margin found in many species of Herklotsichthys and Sardinella : Die Spitzen der Schwanz- flossen lappen und der Innenrand derselben sind dunkel angeftogen. Herklotsichthys punctatus is one of the commonest Indo-Pacific clupeids and Steindachner's descrip- tion fits the species. 8 P. J. P. WHITEHEAD SARDINELLA Valenciennes, 1847 Sardinella Valenciennes, 1847, Hist. Nat. Poiss., 20 : 261 (Type : Sardinella aurita Valenciennes). As yet, there is no modern world-wide study of the Sardinella species with 9 pelvic finrays (5. aurita, S. longiceps, S. anchovia, S. brasiliensis). In the Indo- Pacific region, 5. aurita and 5. longiceps (whose ranges apparently do not overlap) can be easily distinguished (Chan, 1965), but studies of aurita-like fishes in the Atlantic have not compared Eastern with Western Atlantic forms, or both with the Western Pacific 5. aurita, at least on the basis of adequate material. 3. Clupea brasiliensis Steindachner, 1879 = Sardinella brasiliensis (Steindachner, 1879) Clupea brasiliensis Steindachner, 1879, Sitzb. K. Akad. Wiss. Wien, 80 : 182 (fish market, Rio de Janeiro) (non Clupea brasiliensis Schneider, 1801 = Albula vulpes — the Schneider name now a nomen oblitum) ; Idem, 1880, Ichthyol. Beitr., 8 : 64. Clupea Janeiro Eigenmann & Bray, 1894, Ann. N.Y. Acad. nat. Sci. : 626 (replacement name for the preoccupied Clupea brasiliensis Steindachner). SYNONYMY. Clupea brasiliensis Steindachner (a species of Sardinella) is a primary homonym of Clupea brasiliensis Schneider ( = Albula vulpes — Whitehead, 1969^. Fowler (1941 : 602) avoided the issue by resurrecting the earlier Rafinesque name allecia for the Sardinella species, but this name is both doubtfully legal (Whitehead, I967a : 40) and applies to the Mediterranean species of Sardinella ; as shown below, Steindachner's Brazilian species is possibly distinct. Myers (in Rivas, 1964 : 410) believed that a new name was required for the latter, but overlooked Clupea Janeiro of Eigenmann and Bray. The Schneider name has not been used, however, as a senior synonym for over fifty years (see full synonymies in Fowler, 1941 and Hilde- brand, 1964) and it thus qualifies as a nomen oblitum under Article 23 (b) of the International Code. Application will be made to place the Schneider name on the Official Index and thus to release the Steindachner name for this species of Sardinella. TYPE MATERIAL a. LECTOTYPE, a fish of 143-3 mm S.L., ex Rio de Janeiro in 1877, NMV.U56 (jar labelled VII 126 n.sp.). b. PARALECTOTYPES, 2 fishes, 121-0-121-2 mm S.L., ex Rio de Janeiro in 1874, NMV.H58 (jar labelled I 13 . . .). c. PARALECTOTYPES, 2 fishes, 114-4-140-7 mm S.L., ex Rio de Janeiro in 1874, NM V.i 159 (jar labelled I 1356 pt.b). d. PARALECTOTYPES, 4 fishes, 116-7-120-7 mm S.L., ex Rio de Janeiro in 1874, NMV.n6o (jar labelled I I2i6a). e. PARALECTOTYPE, i fish, 120-5 mm S.L., ex Rio de Janeiro, in 1874, NMV. 1161 (jar labelled I 1623). f. PARALECTOTYPES, 4 fishes, 114-0-121-7 mm S.L., ex Rio de Janeiro in 1874, NMV.ii.62 (jar labelled I 1216 pt). STEINDACHNER'S CLUPEOID FISHES 9 g. ? PARALECTOTYPE, i fish, 148-3 mm S.L., ex Rio de Janeiro in 1879, NMV- 1155 (jar labelled coll. Pape 62). This fish may not have been available when the original description was made. DESCRIPTION. A fish, 143-3 mm S.L., 180 mm tot.l. (estimated, caudal tips damaged), LECTOTYPE, e% Rio de Janeiro fish market in 1874, in good condition except caudal, NMV. 1156. Br.St. 6, D v 14, P i 15, V i 8, A iii 15, g.r. 155, scutes 19+14. In percentages of standard length : body depth 22-5, body width 12-4, head length 27-7 ; snout length 7-5, eye diameter 6-6, length of upper jaw n-o, length of lower jaw 13-9, height of lower jaw 6-0, sub-ocular depth 7-3, least post-orbital distance 11-9 ; pectoral fin length 16-9, pelvic fin length 9-7, length of anal base 12-8 ; pre- dorsal distance 45-5, pre-pelvic distance 52-7, pre-anal distance 79-2. Body a little compressed, its width if in depth, belly rounded before pelvic base but scutes more keeled behind ; head length greater than body depth, post-orbital portion a little less than \ length of head. Snout a little longer than eye diameter. Upper jaw toothless, reaching to vertical from eye centre ; two supra-maxillae, the posterior with slender anterior shaft and lozenge-shaped expanded portion posteriorly, its upper and lower profiles meeting anterior shaft at the same point. Lower jaw toothless, rising steeply anteriorly, its height 2\ times in its length, its articulation slightly behind vertical from eye centre. Fine granular teeth present on tongue, palatines and ectopterygoids. Posterior border of operculum with slight indentation in its upper part, lower border horizontal ; sub-operculum rectangular but posterior angle evenly rounded ; inter- operculum about ^ eye diameter at widest point. Posterior border of gill opening with two fleshy lobes ; cleithral lobe present ; isthmus slender, tapering, the sterno- hyoideus muscle extending forward to gill membrane. Gillrakers fine and slender, the longest 8-9 mm (a little more than eye diameter) ; mediopharyngobranchial pre- sent, its length f eye diameter, bearing the first thirty upper gillrakers ; over one hundred slender gillrakers present on posterior face of 3rd epibranchial ; gill filaments of anterior hemibranch on first arch | eye diameter and f length of those of posterior hemibranch. Pseudobranch present, exposed, extending onto inner face of oper- culum, equal to eye diameter, with 20 filaments ; ventral border forming a distinct ridge with a longitudinal groove below for reception of tips of hypobranchial rakers of first arch. Fronto-parietal region of head with two cuneiform areas bearing about ten longitudinal striae, a well-defined transverse ridge on the supra-occipital and several smaller striae above the eyes. Dorsal fin origin nearer to snout tip than to caudal base by if eye diameters ; base of fin invested in low scaly sheath. Pectoral fin tips fail to reach pelvic base by i^ eye diameters ; no axillary scale but a groove above first unbranched ray for reception of fin. Pelvic fin base nearer to pectoral base than to anal origin ; axillary scale present, f length of fin. Anal fin origin nearer to caudal base than to pelvic base by \ eye diameter ; base of fin invested in low scaly sheath, final anal ray much branched and about twice length of antepenultimate ray. Unexposed portion of scales with one continuous and 2 (anterior) to 4 (posterior) io P. J. P. WH1TEHEAD minor vertical striae, the later broken in centre of scale (except for posterior striation in some post-dorsal scales) ; exposed border of scale with fine crenulations and faint horizontal lines. Colour : upper £ of body brown, remainder silvery ; small dark semicircular area on posterior border of operculum due to absence of guanine layer on inner face ; faint dark tips to dorsal branched rays. IDENTIFICATION. Regan (1917 : 378) placed Clupea brasiliensis Steindachner in the synonymy of the widespread Sardinella aurita Valenciennes, but Longley & Hildebrand (1941) noticed higher gillraker counts in three out of four syntypes of Sardinella anchovia Valenciennes ( = S. aurita} and recognized these three specimens as distinct and most likely Steindachner's brasiliensis. Hildebrand (1964 : 399) separated S. brasiliensis from the two other Western Atlantic species, chiefly on the basis of its higher gillraker count (numbers increase in larger fishes, however) , which he cited as : S. brasiliensis 110-130 (160-178 in Longley & Hildebrand, 1941) 5. pinnula Bean 75-80 5. anchovia 55-100 The three doubtful syntypes of S. anchovia (104-5-143-6 mm S.L.) have been re- examined (Whitehead, ig67a : 42) and their high gillraker count was confirmed (151, 174, 179). Another difference between these brasiliensis-like syntypes and the remaining true lectotype of 5. anchovia was found to be the greater depth of the cheek in the former (greater than vertical eye diameter). On the basis of gillraker counts and cheek depth it was accepted that S. brasiliensis differed from other Western Atlantic species and probably also from 5. aurita of the Mediterranean, Eastern Atlantic and Pacific (which is also most likely one of the Western Atlantic species, i.e. the S. anchovia of authors — see Whitehead, ig67a : 43). Examination of specimens of 5. aurita from the Mediterranean, West Africa and the Philippines has now shown, however, that S. brasiliensis cannot be separated from them on its apparently deeper cheek. Furthermore, the total range in gillraker numbers (66-166 fide Chan, 1965 ; up to 280 in 350 mm fishes fide Rossignol, 1955) means that separation will depend on a formula relating gillraker counts to length of fish. Lima (1966) described as S. brasiliensis six specimens of 128-160 mm S.L. from Estado do Ceara, Brazil. Her gillraker counts of 64-128, however, appear to be too low for fishes of this size, at least as far as the figures for the types of brasiliensis are concerned ; on the other hand, they are close to the numbers cited by Hilde- brand (1964). The identify of 5. brasiliensis must depend, therefore, on examination of large samples, especially from the Western Atlantic. Descriptions of the types of brasiliensis, anchovia and aurita reveal only small differences which, although not totally vindicating Regan's synonymizing of the three, at least suggest that no Western Atlantic study can afford to ignore the Eastern Atlantic, Mediterranean and Pacific forms of 5. aurita. STEINDACHNER'S CLUPEOID FISHES n ESCUALOSA Whitley, 1940 Escualosa Whitley, 1940. Aust. Zool., 9 (4) : 402 (Type : Clupea macrolepis Steindachner = Kow- ala thoracata Valenciennes). Generic and species synonymies are discussed in Whitehead (ig64a ; ig67a : 70) and Whitehead et al. (1966 : 70) and a key to this section of the Clupeinae appears in Whitehead (1968). 4. Clupea macrolepis Steindachner, 1879 = Escualosa thoracata (Valenciennes, 1847) Kowala thoracata Valenciennes, 1847, Hist. Nat. Poiss., 20 : 363 (Pondicherry ; type redescribed in Whitehead, ig67a). Clupea macrolepis Steindachner, 1879, Denkschr. A had. Wiss. Wien, 41 (2) : 13 (Townsville, Queensland ; type of Escualosa Whitley, 1940). TYPE MATERIAL. HOLOTYPE, a fish of 64 mm S.L., ex Townsville, Cleveland Bay, Queensland, Australia, coll. Baron Ferdinand von Miiller, SMNS.2292. DESCRIPTION. Proportional and meristic characters for the holotype were given in Whitehead (i964a : 44, Table III). In all respects the type conforms to the more detailed description of the species given in the same publication (p. 45) and in White- head (i967a : 71). SYNONYMY. Clupea macrolepis Steindachner has been shown to be a junior syno- nym of Kowala thoracata Valenciennes (Whitehead, ig64a : 43). Whitley (1940 : 402) designated Clupea macrolepis type of Escualosa Whitley. Since Leptogaster Bleeker proved to be a nomen oblitum (Whitehead et alii, 1966 : 70) and Kowala Valenciennes a synonym oiSardinella (Whitehead, I964a : 52), the genus Escualosa was recognized, containing the single species E. thoracata. Whitley (1940 : fig. 9) figured a specimen of Harengula sp. and suggested that it represented the adult of his Escualosa macro- lepis. The following year (Whitley, 1941 : i) he found in Paris a replacement speci- men of the type of Harengula abbreviata Valenciennes (the real type being missing), identifying it as Kowala castelnaui Ogilby " whose name is obviously a synonym of abbreviata which belongs to my genus Escualosa, 1940 ". In fact, the replacement specimen does not match the description of abbreviata and the latter name should be considered a nomen dubium (Whitehead, i^6ya. : 69). Whitley's specimen of Haren- gula appears to be Herklotsichthys castelnaui and Escualosa remains a monotypic genus distinguished by possession of 7 pelvic rays, a silver lateral stripe and a large rectangu- lar 2nd supra-maxilla (see key, Whitehead, 1968 : 478). RHINOSARDINIA Eigenmann, 1912 Heringia Fowler, 1911, Proc. Acad. nat. Sci. Philad., 63 : 207 (Type : Clupea amazonica Stein- dachner) (name preoccupied in Diptera — Myers, 1929, Copeia : i). Rhinosardinia Eigenmann, 1912, Mem. Carnegie Mus., 5 : 445 (Type : Rhinosardinia serrata Eigenmann = R. amazonica Steindachner). The curious retrorse spine at the anterior end of the maxilla sets this genus apart from all others (see key in Whitehead, 1968 : 478). A small projection occurs in 12 P. J. P. WHITEHEAD Escualosa (Whitehead, ig6^a. : fig. 20) but this is blunt and points upwards and not backwards. In other respects Rhinosardinia is very similar to the South American genera Lite and Ramnogaster. Regan (1917 : 394) placed Rhinosardinia (as Heringia) with Lile, Sardinella and Harengula, i.e. in the Clupeinae. Hildebrand (1964 : 261) allied Rhinosardinia with the Pristigasterinae, but current definitions of the clupeid subfamilies (e.g. Whitehead et alii, 1966 : 37), although by no means satisfactory, exclude Rhinosardinia from the Pristigasterinae, because of its short anal fin (under 20 rays ; cf. over 30), and place it in the Clupeinae. At species level, the South American clupeids show rather little relationship to the West African or Indo-Pacific clupeids (only Sardinella aurita is found on both sides of the Atlantic), and only the genera Sardinella, Ilisha and Pellona are shared (the last not from West Africa). It has been argued, therefore (Whitehead, 1968), that the South American Clupeinae have evolved in isolation. The similarities between Rhinosardinia and the Indo-Pacific Escualosa may thus be coincidental, resulting from relatively limited variations possible within the clupeine framework. In Escualosa, as also in Rhinosardinia (but less so in R. bahiensis) the upper jaw is more or less notched, a condition typical of the shads (subfamily Alosinae) . Although Brevoortia, Ethmidium, Ethmalosa and the Indo-Pacific shads (Hilsa, Gudusia) have very long and numerous gillrakers and no jaw teeth, teeth and short, sparse rakers are found in some species of Alosa (sensu Svetovidov, 1964, i.e. including Pomolobus and Caspialosa). Future work may show that Rhinosardinia can be derived as con- vincingly from the Alosa stem as from the Western Atlantic Clupeinae. 5. Clupea amazonica Steindachner, 1879 = Rhinosardinia amazonica (Steindachner, 1879) Clupea amazonica Steindachner, 1879, Sitzb. K. Akad. Wiss. Wien, 80 : 183 (Amazon R. at Para ; 4 fishes, 40-75 mm tot. 1.) ; Idem, 1880, Ichthyol. Beitr., No. 8 : 65 (repeat). Rhinosardinia serrata Eigenmann, 1912, Mem. Carnegie Mus., 5 : 445, text fig. 39, pi. 62 (figs. 3 and 4) (ex Morawhanna and Mora Passage, British Guiana). TYPE MATERIAL, a. LECTOTYPE, a fish of 54-8 mm S.L. (74-0 mm tot.l.), ex Para, Amazon River in 1879, NMV.HO4. b. PARALECTOTYPES, 3 fishes 31-2-42-2 mm S.L. from the same jar. Steindachner listed only 4 fishes. A fifth specimen in this jar (25-9 mm S.L., 32 mm tot.l.) is too small to have been part of the original description. DESCRIPTION. A fish 54-8 mm S.L. (74-0 mm tot.l.) ex Para, Amazon River, in good condition, NMV.H04. Br.St. 5 (left 6), D ii 13, P i n, V i 7 (both sides), A iii 13, g.r. 38, scutes 17+11. In percentages of standard length : body depth 30-3, head length 24-1 ; snout length 5-75, eye diameter 7-3, length of upper jaw 9-85, length of lower jaw 9-5, depth of lower jaw 5-6 ; pectoral fin length 19-8, pelvic fin length 12-4, length of anal fin base 13-0, length of dorsal base 14-2, height of dorsal fin 18-6 ; pre-dorsal distance 51-0, pre-pelvic distance 51-0, pre-anal distance 76-5 ; depth of caudal peduncle 13-3. Body compressed, belly with trenchant keel of scutes, body depth greater than head length. Snout shorter than eye diameter. Upper j aw with two supra-maxillae, STEINDACHNER'S CLUPEOID FISHES the ist (anterior) slender, the 2nd (posterior) with diamond shaped expanded portion reaching to posterior tip of maxilla ; minute teeth present on lower edge of maxilla below centre of 2nd supra-maxilla, and a sharp retrorse spine ( = to half pupil diameter) on upper edge of maxilla in front of anterior supra-maxilla ; upper jaw with a slight median notch. Lower jaw profile rising steeply, its deepest part in first £ of its length, the jaw 1-6 times as long as deep. No teeth on pre-maxillae or lower jaw or within mouth. Gillrakers fine and slender, £ eye diameter and a little longer than corresponding gill filaments ; about 10 short rakers on posterior face of 3rd epibranchial. Pseudo- branch f eye diameter, with n filaments. Operculum with anterior and posterior margins parallel, the lower border rising at about 40° ; sub-operculum rectangular with rounded posterior border ; inter-operculum deep posteriorly, tapering evenly to the lower jaw articulation. Fronto-parietal region of head smooth, posterior frontal fontanelles occluded. Dorsal fin origin midway between tip of snout and base of caudal fin ; dorsal height (last unbranched ray) 1-30 times length of dorsal base. Pectoral fin failing to reach pelvic base by just over i eye diameter ; no axillary scale present. Pelvic fin base in advance of dorsal origin by almost I pupil diameter and equidistant between pectoral base and anal origin ; axillary scale present, ^ length of fin. Anal origin equidistant between pelvic base and base of caudal fin. Depth of caudal peduncle 2-27 times in body depth. Unexposed portion of scales with one continuous vertical striation and up to three small radiating striae not reaching to centre of scale ; exposed portion of scale with two horizontal striae converging to centre of scale. Small scales present on caudal. Colour : overall brownish, but with no suggestion of a silver lateral stripe (absent also in paralectotypes) . SYNONYMY. Regan (1917 : 394) considered R. serrata Eigenmann a synonym of R. amazonica, but Hildebrand (1964 : 415) after " careful rereading of the original description of amazonica " decided to separate the two on small differences in body S.L. As % of S.L. Body depth Head length Caudal peduncle depth Dorsal fin : base height height base Body depth Caudal ped. 1. Gillrakers R. amazonica (LECTOTYPE) NMV.H04 54-8 30-3 24-1 13-3 14-2 18-6 2-27 38 R. serrata (SYNTYPE) USNM.66284 47-3 30-0 23-8 13-1 12-7 18-0 1-42 2-29 35 R. serrata (4 SYNTYPES) BMNH.igii.io. 3I-452-5 44.5-49-6 28-5-30-5 21-6-23-3 12-7-13-5 12-2-13-8 18-7-20-0 i -43-1 -49 2-18-2-31 33. 34. 34. 35 i4 P. J- P. WHITEHEAD depth, depth of caudal peduncle, head and snout length and length of dorsal fin base. A comparison between five syntypes of R. serrata and the lectotype of R. amazonica does not bear out Hildebrand's distinction between these two nominal species. The slightly longer dorsal fin base in the lectotype of R. amazonica and its slightly higher gillraker count hardly justify separation of the two species. The range of R. amazonica thus extends from the mouth of the Amazon, through the Guianas to the San Juan river in Venezuela. 6. Pellonula bahiensis Steindachner, 1879 = Rhinosardinia bahiensis (Steindachner, 1879) (Plate ib) Pellonula bahiensis Steindachner, 1879, Sitzb. K. Akad. Wiss. Wien, 80 : 181, pi. 3 (fig. 2) (ex Bahia, 8 fishes to 100 mm tot.l.) ; Idem, 1880, Ichthyol. Beitr., No. 8 : 63, pi. 3 (fig. 2) (repeat). TYPE MATERIAL. LECTOTYPE, a fish of 69-1 mm S.L. (90-5 mm tot.l.), in good condition but some scales missing, ex Bahia in 1879, NMV.287O. PARALECTOTYPE, a fish of 70-5 mm S.L. from the same bottle. DESCRIPTION. A fish, 69-1 mm S.L. (90-5 mm tot.l.), LECTOTYPE, ex Bahia, NMV.2870. Br.St. 6 (6 right), D iii 15, P i 13, V i 7, A iii 14, g.r. 32, scutes 17+11. In percentages of standard length : body depth 27-3, body width II-T, head length 25-0 ; snout length 6-4, eye diameter 8-7, length of upper jaw 11-9, length of lower jaw 11-9, depth of lower jaw 6-2 ; pectoral fin length 15-9 pelvic fin length 12-7, length of dorsal fin base 17-5, height of dorsal fin 19-0, length of anal fin base 16-2 ; pre-dorsal distance 46-1, pre-pelvic distance 49-8, pre-anal distance 72-5. Body compressed, its width 2| times in its depth, belly keeled, scutes with long sharp spines partly concealed by scales on either side of midline ; head a little shorter than body depth. Snout a little shorter than eye diameter. Upper jaw reaching almost to vertical from eye centre ; two supra-maxillae, the ist (anterior) long and plate-like, the 2nd (posterior) of typical Harengula shape and reaching to posterior tip of jaw ; maxilla with a small blunt projection (? damaged ; — retrorse spine in paralectotype) on upper face of maxilla in front of ist supra-maxilla ; minute denticulations along lower edge of maxilla below 2nd supra-maxilla ; upper jaw without marked notch. Lower jaw profile rising steeply, its deepest part in first ^ of its length, the jaw 1-91 times as long as deep. Small conical teeth present at dentary symphysis (left 4, right 3), on pre-maxillae (left 4, right 7) and even smaller on palatines and ecto-pterygoids. Gillrakers fine and slender, the longest just under half eye diameter, i£ times length of corresponding gill filaments ; seven short triangular rakers present on pos- terior face of third epibranchial ; short mediopharyngobranchial present, bearing 6 gillrakers. Pseudobranch present, exposed, ^ eye diameter, with 14-15 filaments. Cleithral lobe present. Exposed portion of inter-operculum a narrow crescent, less than half depth of sub-operculum. Frontals diverging in the midline posteriorly to expose a triangle of the supra-occipital ; posterior frontal fontanelles not exposed ; STEINDACHNER'S CLUPEOID FISHES 15 fronto-parietal region with a series of short longitudinal striae (not as strongly developed, however, as in e.g. Sardinella). Dorsal fin origin nearer to snout than to caudal by just over i eye diameter and in advance of vertical from pelvic base by i pupil diameter ; base of fin 1-09 times in its height. Pectoral fin tips failing to reach pelvic base by just over i eye diameter ; axillary scale absent (apparently) but a short groove present above proximal £ of first ray. Pelvic fin base nearer to pectoral base than to anal origin by | pupil diameter ; axillary scale present, a little over \ length of fin. Anal fin origin about equidistant between pelvic and caudal bases ; a low sheath of scales along base of fin. Scales : unexposed portion of scale with one major and up to three minor striae, the former vertical and continuous across scale, the latter radial and only occasionally traversing scale ; exposed portion of scale without striae or with one or two very short radial striae. Colour : body brown, with very distinct silver lateral band from opercular opening to caudal base, almost as broad as eye ; opercular series and belly silvery. NOTE. Regan (1917 : 395) doubted that Pellonula bahiensis could be separated from Clupea amazonica on the basis of its silver lateral stripe, since specimens of the latter " also show a faint lateral band in certain lights. " The stripe in R. bahiensis is, however, quite as distinct as that which separates e.g. Spratelloides gracilis from 5. delicatulus in all except post-larval size groups. It was well illustrated by Stein- dachner (see Plate ib). Rhinosardinia amazonica and R. bahiensis were clearly distinguished by Hilde- brand (1964 : 411), but examination of the types necessitates some modifications to the key (as stated earlier, R. serrata is a synonym of R. amazonica). KEY TO SPECIES Rhinosardinia 1 Silver lateral stripe present ; exposed portion of scales without 2 prominent horizontal striae ; dorsal fin origin nearer to snout than to caudal base by i eye diameter ; inter-operculum visible as narrow crescent, £ as deep as sub-operculum . . Rhinosardinia bahiensis (Steind.) 2 No silver lateral stripe ; exposed portion of scales with 2 prominent horizontal striae ; dorsal fin origin equidistant between snout tip and caudal base ; inter-operculum broadly exposed, at its widest as deep as sub-operculum . . . Rhinosardinia amazonica (Steind.) SARDINOPS Hubbs, 1929 Sardinops Hubbs, 1929, Proc. Calif. Acad. Sci., 18 (n) : 264 (Type : Meletta caerulea Girard). As in other wide-ranging genera with a discontinuous distribution, the various forms of Sardinops have sometimes been considered to be distinct species and some- times merely subspecies. Regan (1916 : 14) and Chabanaud (1926) suspected the latter but excluded S. neopilchardus of Australia and New Zealand ; Svetovidov (1952 : 177) recognized five subspecies of Sardinops sagax, but with some reservation regarding neopilchardus, in which the head is longer, the maxilla shorter and the gillrakers less numerous than in the other forms. On present evidence, Svetovidov's solution seems best. 16 P. J. P. WHITEHEAD 7. Alausa fimbriata Kner & Steindachner, 1866 = Sardinops sagax sagax (Jenyns, 1842) (Plate ic) Clupea sagax Jenyns, 1842, Zoo/. Beagle, Fish. : 134 (ex San Lorenzo I., Lima). Alausa fimbriata Kner & Steindachner, 1866, Sitzb. K. Akad. Wiss. Wien, 54 : 386, pi. 15 (" Val- paraiso? ", no size, from Museum Godeffroy). LOCALITY. As noted earlier (p. 6), Steindachner confused the reference number and locality of this species with that of his Alausa alburnus. The correct locality is thus Valparaiso, Chile. TYPE MATERIAL. Intensive search produced no specimen labelled Alausa fim- briata in Vienna. But Steindachner's description and figure are quite sufficient to identify this species and provision of a neotype would serve no useful purpose. IDENTIFICATION. Steindachner's pi. 15 (shown here, Plate ic) shows the striations on the operculum, the enlarged anal rays and the large alar scales that characterize members of Sardinops ; the description is quite consistent with this identification. A series of black spots often occurs along the flank but this was apparently not the case in Steindachner's specimen. Steindachner appears to have been unaware of Jenyns work on the Beagle collec- tion. 8. Clupea neopilchardus Steindachner, 1879 = Sardinops sagax neopilchardus (Steindachner, 1879) Clupea lata Richardson, 1843, Trav. N.Z. (Dieffenbach), 2 : 221 (on Solander MS. name Clupea lata — nomen nudum). Clupea neopilchardus Steindachner, 1879, Denkschr. Akad. Wiss. Wien, 41 : 12 (not quite 170 mm, ex Hobson's Bay, Victoria, type in Stuttgart Museum). TYPE MATERIAL. The type is apparently no longer present in Stuttgart. This sub-species is sufficiently well defined geographically (southern Australia and New Zealand — nearest related forms in South Africa and along Pacific coast of South America) for the provision of a neotype to be unnecessary at the present time, particularly in view of the excellent study of the Australian form by Blackburn (1949). Richardson's Clupea lata is not accompanied by any description but merely a reference to Solander's MS. description in his notebook (Pisces Auslraliae, p. 17) made during Captain Cook's first voyage to the Pacific. Solander's description reads : Clupea lata B.i8 Habitat Tolaga Argentea, nitidipinna ; Dorso e cinereo caeruleo nitente, ut et superna pars capitis. Pinna analis basi plumbea, alias omnes colore corporis ubi sita. Pinna caudalis tota plumbea Iris argentea, superne semper nebula nigra. STEINDACHNER'S CLUPEOID FISHES 17 This description is not adequate to identify the species and no drawing of Clupea lata was made by Sydney Parkinson, the artist on this voyage. The name lata is a nomen nudum. 9. Clupea setosa Steindachner, 1869 = Ethmalosa fimbriata (Bowdich, 1825) (Plate 2a) Clupea fimbriata Bowdich, 1825, Excurs. Madeira : 234, fig. 44 (Porto Praya, Cape Verde Is. — probably the Gambia fide Whitehead, i96yb : 590). Clupea setosa Steindachner, 1869, Sitzb. K. Akad. Wiss. Wien, 60 : 311, pi. 6 (ex Mazatlan stated — in fact West Africa) ; Idem, 1869, Ichthyol. Notizen, No. 9 : 22, pi. 6 (repeat). LOCALITY. Steindachner (1882 : 14) subsequently realized that his specimens had not come from Mazatlan (Mexico) but were from the coasts of Liberia and Gabon, West Africa. TYPE MATERIAL. LECTOTYPE, a fish of 184-0 mm S.L. (253-0 mm tot.l.), ex West Africa (Liberia or Gabon coasts), NMV.4I73. PARALECTOTYPES, two fishes, 174-5-178-0 mm S.L., as above, caudal lobes damaged. The specimens are accompanied by labels reading : " Coll. Musei Vindobonensis 4173 CLUPEA altata Std. ALTATA Steind. 1878 I 20 " and " Coll. Musei Vindobonensis 4173 CLUPEA setosa Steind. ALTATA (Mexico Sinaloa) 1878 ". DESCRIPTION. A fish, 184-0 mm S.L. (253-0 mm tot.l.), LECTOTYPE, ex West Coast of Africa, in good condition. NMV.4I73 Br.St. 6, D iv 14, P i 14, V i 7, A iii 18, g.r. 130, scutes 18+12. In percentages of standard length : body depth 39-4, body width 13-4, head length 34-0 ; snout length 8-3, eye diameter 7-5 ; upper jaw length 15-5, lower jaw length 18-0, post-orbital distance 18-7 ; pectoral fin length 21-8, pelvic fin length 12-7, length of anal fin base 17-9 ; pre-dorsal distance 49-2, pre-pelvic distance 56-2, pre-anal distance 76-5- Body compressed, its width 3 times in its depth, the latter a little greater than head length ; belly keeled, but scutes lying in narrow groove formed by scales and thus scarcely projecting below profile of body. Head with adipose tissue covering much of upper part, adipose eyelid present and covering all but a third of eye. Cutaneous sensory canals on operculum, pre-operculum and 2nd sub-orbital and continued on scales behind occiput and around upper border of gill opening. Snout a little longer than eye. Upper jaw with distinct median notch into which distal tip of lower jaw fits. Maxilla without longitudinal ridges, reaching posteriorly to vertical from just behind eye centre ; two supra-maxillae. No teeth in jaws or on tongue. Operculum with lower border rising steeply upwards ; anterior border of operculum with slight cut-away, exposing junction of sub- and inter-opercula. i8 P. J. P. WHITEHEAD Pseudobranch present, exposed, a little greater than snout length, the lower border forming a distinct ridge with a groove below it. Gillrakers as described in the neotype of Clupea fimbriata Bowdich (see Whitehead, igbyb : 591) ; longest rakers on lower arm of first arch equal to eye diameter and about three times length of cor- responding filaments. Dorsal fin origin nearer to snout than to caudal base by f eye diameter ; first un- branched ray very small. Pectoral fin tip almost reaching to pelvic base ; one large and two small scales in axil of fin, forming a groove along £ length of fin. Pelvic fin base nearer to pectoral base than to anal origin by £ eye diameter. Anal fin origin nearer to pelvic base than to caudal base by \ eye diameter. Scales adherent, about 40 in lateral series ; posterior edges fimbriated, as in Steindachner's figure (see Plate 2a). Colour : uniform brown with dark brown at tips of anterior dorsal rays. IDENTIFICATION. The genus Ethmalosa is monotypic and occurs only off the West Coast of Africa. The synonymy of the species is fully dealt with by Whitehead (1967^. The relationship of Ethmalosa to other genera of shads is discussed by Whitehead (i965b : 153). 10. Clupea notacanthoides Steindachner, 1869 = Ethmidium maculatum notacanthoides (Steindachner, 1869) (Plate 2 b) Alausa maculata Valenciennes, 1847, Hist. Nat. Poiss., 20 : 430 (Valparaiso — holotype redes- cribed by Whitehead, ig6ja, : 88). Clupea notacanthoides Steindachner, 1869, Sitzb. K. Akad. Wiss. Wien, 60 : 309, pi. 7 (Mazatlan stated — probably erroneous) ; Idem, 1869, Ichthyol. Notizen, No. 9 : 20, pi. 7 (repeat). Ethmidium chilcae Hildebrand, 1946, Bull. U.S. natl. Mus., No. 189 : 82 (Callao and Chilka Bay, Peru). LOCALITY. No mention of Steindachner's species is made by Jordan (1895) in his list of fishes from Sinaloa, nor in the check-list of Jordan, Evermann & Clark (1930). In view of the mislabelling of the ' Mazatlan ' specimens of Clupea setosa (see p. 17), it seems probable that Steindachner's Clupea notacanthoides came from the normal range of Ethmidium maculatum, i.e. Peru or Chile. TYPE MATERIAL. Intensive search in Vienna has failed to produce the specimen (s) on which the description was made ; no size is stated, but the figure suggests a fish of about 175 mm S.L. (see Plate 2b). DESCRIPTION, (based on Steindachner's description). Br.St. 9, D 20, P 17, V 7, A 14, scales in lateral series ca 50, 7 predorsal scutes (? error for 17), ventral scutes ca 18+17. In percentages of standard length : body depth 30, head length 32. In percentages of head length : snout length 21, eye diameter 15-8, interorbital 25, lower jaw 55-5 ; longest dorsal ray 44-5, pectoral length 36-5 (57 in figure). Dorsal origin if eye diameters nearer to snout than to caudal base. Pectoral falling short of pelvic base by just over i eye diameter (according to figure) ; a series STEINDACHNER'S CLUPEOID FISHES 19 of enlarged scales above first ray. Pelvic base equidistant between tip of snout and caudal base. Anal base about one eye diameter shorter than head length ; longest ray i^ eye diameters. Colour : a series of six black spots on the flank below the midlateral line. IDENTIFICATION. The well illustrated dorsal scutes and high branchiostegal count confirm that Steindachner's specimen was a species of Ethmidium. Hildebrand (1946 : 84-85) proposed a new species, Ethmidium chilcae, for his Peruvian specimens, which he distinguished from E. maculatum of Chile in the following way. E. maculatum E. chilcae Lota, Chile Callao & Chilka Bay, Peru (70-113 mm S.L.) (ca 100-212 mm S.L.) Head in S.L. 3'25-3'4 (29-4-30-8%) 3-0-3-1 (32-3-33-3%) Depth in S.L. 2-75-2-9 (34-5-36-4%) 2-8-3-1 (32-3-35-7%) Caudal peduncle depth in head length 2-80-2-95 3-20-3-75 Pectoral fin tip short of pelvic base by eye diam. by pupil diam. or " less than eye " in adults Ventral profile more convex Scales nearly smooth denticulate Examination of larger specimens from Chile (over 200 mm S.L.) shows that in adults the convexity of the belly profile, the denticulations on the posterior edge of the scale and the depth of the body reach the condition specified by Hildebrand for his Peruvian E, chilcae (e.g. in the type of E. maculatum from Valparaiso as redescribed in Whitehead, 19673. and in two British Museum specimens from the Gulf of Arauco, Chile). The only Peruvian specimen in the British Museum collection is a large adult, 245 mm S.L., which agrees with Hildebrand's diagnosis of E. chilcae in having a large head (35-8% of S.L.), a caudal peduncle depth 3-65 times in head length and denticulate scales. Unfortunately, the belly and pelvic fins are missing. Eight Peruvian specimens in the Copenhagen collections (ex Callao, 180-266 mm S.L., ZMC. 544-5 and 18289-91) agree with Hildebrand's diagnosis in head length (31-1-33-0% of S.L.), body depth (327-36-2% of S.L.), caudal peduncle depth in head length (3'°7-3'75 times) and distance between pectoral tip and pelvic base (i£-2i pupil diameters, but less than eye). The scales bear 20-22 denticulations on the posterior margin. On the basis of the material examined and the small size of Hildebrand's Chilean specimens, it is likely that head length may distinguish Peruvian from Chilean stocks, but that body depth, scale form and apparent pectoral length are probably dependent on the size of the fish or on exogenous factors (trophic conditions). Mann (1954) regarded Hildebrand's Peruvian material as a subspecies of E. maculatum and stated that it reached as far south as Antofagasta in Chile, its place then being taken by the nominate form. Two small British Museum specimens (82-92 mm S.L.) from Herradura Bay, just north of Antofagasta, clearly fit Hilde- brand's diagnosis of E. maculatum. For the present, the two forms will be considered subspecies. 20 P. J. P. WHITEHEAD SYNONYMY. Hildebrand (loc. cit.} allied his E. chilcae most nearly to Stein- dachner's Clupea notacanthoides, but noticed slight differences in some proportions. Since it seems very likely that Steindachner had a Peruvian fish — and the description and figure suggest the large-headed Peruvian form— the slight differences noted by Hildebrand may well be attributable to faults in Steindachner's description, in which case the name notacanthoides should be used for the Peruvian subspecies. Gunther's type of Clupea notacanthus from Valparaiso (89-0 mm S.L., BMNH. 1848.6.14.42 — one specimen now missing) is clearly the nominate form (head 30-6% of S.L.). ILISHA Richardson, 1846 Ilisha Richardson, 1846, Kept. Ichthyol. China Japan : 306 (Type : Ilisha abnormis Richardson = Alosa elongata Bennett fide Whitehead, 1966). Platygaster Swainson, 1838, Nat. Hist. Animals, 1 : 278 (Type : Clupea africana Bloch, desig- nated by Swain, 1882, Proc. Acad. nat. Sci. Philad. : 280) ; ibidem., 1839, 2 : 186, 294 (name preoccupied in Hymenoptera). Zunasia Jordan & Metz, 1913, Mem. Carnegie Mus., 6 (i) : 7 (Type : Pristigaster chinensis Basilewski = Ilisha elongata, see below). Pseudochirocentrodon Miranda-Ribeiro 1923, Comm. Linhas Telegr. Estrat. Matto Grosso A mazonas, 58 : 8 (Type : P. amazonicum Mirando-Ribeiro) . Euplatygaster Fowler, 1934, Proc. Acad. nat. Sci. Philad., 85 : 246 (Type : Pellona brachysoma Bleeker = Ilisha indica, see below). At present, the genera Ilisha and Pellona are separated solely on the presence of a toothed hypo-maxilla in the latter (replacing the ligament connecting the tip of the pre-maxilla to the ventral edge of the maxilla) . The presence of a toothed hypo- maxilla also separates the New World Harengula from the Indo-Pacific Herklotsich- thys (subfamily Clupeinae) , and the New World Pacific coast Pliosteostoma from the Atlantic coast Odontognathus (subfamily Pristigasterinae) . Having found a specimen of the Ilisha-Pellona complex with the hypo-maxilla present on one side and absent on the other, Myers (1950) preferred to await other evidence before splitting the genus Ilisha. Hildebrand (1964 : 415) also combined Ilisha and Pellona, but Berry (1964 : 729) split them again. No supporting evidence has yet been published, and I have been unable to find consistent differences in gill arches, fronto-parietal striation patterns, shapes of bones in the opercular and maxillary series, scutes or scales. Nevertheless, the separation of the species on a single, easily determined character (the hypo-maxilla) is useful and the generic level is preferred here. It is unfortunate that the separation of Pellona from Ilisha does not coincide with a geographical separation. Most of the Indo-Pacific species are Ilisha, but Pellona ditchella Valenciennes has a hypo-maxilla ; conversely, most of the New World species are Pellona, but two species of Ilisha are recognized. I have been unable to find consistent differences between the groups of species from the two geographical areas. The only comprehensive key to the species of Ilisha is that of Norman (1923). Subsequent keys are also unsatisfactory in view of re-examination of the type STEINDACHNER'S CLUPEOID FISHES 21 material of Richardson, Bleeker, Valenciennes, Bloch and Steindachner (Whitehead, 1966 ; Whitehead et al., 1966 ; Whitehead, igGya; Whitehead, 19690 ; and the present study), coupled with the work of Myers (1950) and Hildebrand (1964). The following key is by no means definitive. It is intended as a summary of recent published and unpublished work ; there is still great need for a full revision of the Indo-Pacific species. KEY TO SPECIES Ilisha A. NEW WORLD 1 Post-pelvic scutes 6-7, total scutes 25-26 ; gillrakers 19-22 ; pelvic base nearer to anal origin than to pectoral base ; Atlantic drainage (Amazon of Brazil, Peru) i. /. amazonica (Miranda- Ribeiro, 1923) 2 Post-pelvic scutes 12-14, total scutes 34-39 ; gillrakers 23-24 (and 11-12 on upper arch) ; pelvic base equidistant between anal origin and pectoral base or slightly nearer to the latter ; Pacific drainage and coasts (Costa Rica to Ecuador) . 2. I.furthii (Steindachner, 1875) B. EASTERN ATLANTIC 3. I. africana (Bloch, 1795) C. INDO-PACIFIC 1 Anal origin in advance of or below middle of dorsal base ; pre-pelvic scutes 24-27 ; gillrakers 17-22 (Burma, Java, Borneo) .... 4. I. pristigastroid.es (Bleeker, 1852) 2 Anal origin below posterior half of dorsal base a. Pre-pelvic scutes 22-27, post-pelvic 10-14 •' gillrakers 20-24 i Body deep, 37% of S.L. ; Borneo . . . 5. I. macrogaster Bleeker, 1866 ii Body depth moderate, 30-34% of S.L. ; India, Borneo ..... 6. I. filigera (Valenciennes, 1847) iii Body slender, 24-28% of S.L. ; India to China . 7. I. elongata (Bennett, 1830) b. Pre-pelvic scutes 18-20, post-pelvic 7-9 i Body depth 37-41% of S.L. ; gillrakers 23-28 ; India to Singapore . 8. I. indica (Swainson, 1839) ii Body depth 32-34% of S.L. ; gillrakers 19-21 ; India to Singapore . 9. I. megaloptera (Swainson, 1839) There appear to be 39 nominal species referable to the genus Ilisha. To summarize recent work, these nominal species are listed (alphabetically) in Table 2, each name preceded by a number which allocates it to a species in the key given above. Follow- ing each entry is a reference (in parenthesis) to works in which type material or synonymies are discussed. Major difficulties surround species groups 5-7 and 8-9, in which body depth is used to separate species. Intraspecific variation in body depth is very poorly documented. The slender I. elongata is fairly distinctive, but the macrogaster-filigera complex may prove to be a single species. Similarly, the indica-megaloptera complex may also comprise a polytypic species in which varia- tions in body depth and gillraker numbers can be correlated with habitat (purely freshwater, marine and intermediates). The Swainson names megalopterus and indicus, based respectively on Jangarloo and Ditchoee of Russell (1803 : pi. 191 and pi. 192 — reproduced in Whitehead, ig67a : pi. 8a, b), must be provided with neotypes. Russell's figure of Jangarloo shows a fish with 17+11 (? 18+12) scutes, a combination not yet found in Indo-Pacific specimens ; the count is presumed to have been 18+9 (i.e. section 2b of the key 22 P. J. P. WHITEHEAD above) . Russell's figure of Ditchoee closely resembles Bleeker's figure and holotype of Ilisha brachysoma (Whitehead, ei al., 1966 : 98, pi. 13 (i) — from figure in Atlas), except that Russell gave an anal count of only 37 (47 rays in the holotype of brachy- soma, but only 40 in the type of Pellona micropus}. Even if Russell miscounted, a variation of 7 rays is rather large and it may later prove necessary to separate /. brachysoma from /. micropus (i.e. /. indica). ii. Pellona furthii Steindachner, 1875 = Ilisha furthii (Steindachner, 1875) Pellona furthii Steindachner, 1875, Sitzb. K. Akad. Wiss. Wien, 70 : 388 (Bay of Panama, to nj zoll in length, i.e. 299 mm) ; Idem, 1875, Ichthyol. Beitr., No. I : 14 (repeat). TYPE MATERIAL a. LECTOTYPE, a fish of 201 mm S.L. (ca 260 mm tot.l. — caudal tips broken) ex Panama Bay in 1874, NMV.mo [jar labelled I 1253 Steind. don. (typ)]. b. PARALECTOTYPE, i fish, 215 mm S.L. (as above). c. PARALECTOTYPES, 2 fishes, 200-217 mm S.L., NMV.no6 (otherwise as above) . d. PARALECTOTYPE, i fish, 210 mm S.L., NMV.im (otherwise as above). e. PARALECTOTYPE, i fish, 212 mm S.L., NMV.I872 (otherwise as above). f. PARALECTOTYPES, 2 fishes, 210-217 mm S.L., NMV.IH4 (otherwise as above). Another fish, 232 mm S.L., ex Panama in 1876 (NMV.i883) is not part of the type series. DESCRIPTION. A fish, 201 mm S.L., 260 mm tot.l. (estimated, caudal tips dam- aged), LECTOTYPE, ex Panama Bay in 1874, in good condition apart from loss of caudal tips, NMV.mo. Br.St. 6, D iv 13, P i 14, V i 5 (both), A iii 45, g.r. 11+23, scutes 22+13. In percentages of standard length : body depth 36-4, body width 9-2, head length 247 ; snout length 6-8, eye diameter 8-0, upper jaw length 13-1, lower jaw length 13-3 ; length of pectoral fin 19-7, length of pelvic fin 6-4, length of anal fin base 45-0 ; pre-dorsal distance 45-7, pre-pelvic distance 43-4, pre-anal distance 60-0. Body strongly compressed, its width 4 times in its depth, the latter i£ times head length ; dorsal profile irregular, with slight " hump " behind occiput, ventral profile evenly convex, scutes prominent especially behind pelvic fin base. Eye large, its diameter greater than snout length and 3 times in head length. Upper jaw reaching to just before vertical from eye centre ; median | of pre-maxillae toothless, a single series of minute conical teeth lateral to this ; no hypo-maxillae ; lower edge of maxillae with a series of fine teeth ; two supra-maxillae present, the ist (anterior) slender and \ eye diameter, the posterior with slender anterior shaft and expanded posterior part reaching almost to tip of maxilla (Figure i). Lower jaw strongly projecting (7-5 mm beyond tip of snout), upper border rising, its height reach- ing half length of jaw just before midpoint ; a single series of 6-10 small conical teeth projecting inwards on either side of the symphysis. STEINDACHNER'S CLUPEOID FISHES 23 Operculum with indentation along posterior border, its lower edge rising fairly steeply. Sub-operculum longer than deep, almost triangular. Entire gill opening covered by opercular series. Pseudobranch present, exposed, a little under £ eye diameter in length, bearing about 25 filaments. Gillrakers slender, the longest £ eye diameter ; gill filaments of anterior and posterior hemibranchs subequal, the longest 2-8 times in eye diameter. Four stubby gillrakers on posterior face of 3rd epibranchial. Granular teeth present on tongue, palatines and endo- and ectopterygoids ; no teeth on vomer. FIG. i. Pellona furthii LECTOTYPE, 201 mm S.L., NMV.mo ( = Ilisha furtMi) . Dorsal fin origin nearer to snout than to caudal base by I eye diameter ; the first unbranched ray short and barely apparent. Pectoral fin tip broken but probably reaching to half-way along pelvic ; axillary scale present, 2^-2^ times in length of fin ; base of fin nearer to snout tip than to pelvic base by £ pupil diameter ; base of fin covered by scales. Pelvic fin base equidistant between pectoral base and anal origin ; no axillary scale present (? lost). Anal origin nearer to caudal base than to snout tip by I eye diameter. Scales partly lost, posterior (exposed) border with faint signs of erosion and some small radiating striae (especially in posterior scales) ; unexposed portion of scales with up to eight convex striae, all interrupted at centre of scale except the most posterior. Colour : upper i/io of body brown, remainder of flanks silvery ; tip of pectorals and tips of posterior dorsal rays speckled brown. 24 P. J. P. WHITEHEAD IDENTIFICATION. The absence of a toothed hypo-maxilla in Pellona furthii was not unexpected. Norman (ig23a) originally placed this species in his genus Neosteus ( = Pellona) implying the presence of this bone, but assigned it to Ilisha when he had examined material (Norman, 1923^. Hildebrand (1946 : 91, footnote) could find no hypo-maxilla in six specimens from the Gulf of Guayaquil and, recognizing Norman's separation of Pellona from Ilisha on this character, placed his specimens in Ilisha. Following the synonymy put forward by Myers (1950 : Ilisha iquitensis and I. apapae synonyms of Pseudochirocentrodon amazonicum), Berry (1964 : 729) deduced that there is a single species of Ilisha in the Western Atlantic drainage area of South America, Ilisha amazonica (Miranda-Ribeiro). No Western Atlantic specimens of Ilisha are in the British Museum collections, but I. amazonica is clearly distinct from /. furthii of Pacific coasts according to the descriptions of Hildebrand (1948 and 1964 : 421) and Myers (1950). The differences are shown in the key given above (p. 21). Scute and some gillraker counts were made on the following seven British Museum specimens and the results incorporated in the key, together with counts for the types of P. panamensis (see below) . a. 4 fishes, 160-201 mm S.L., ex Guayas River, Ecuador, BMNH. 1938. 11.18.1-4. b. 3 fishes, 216-238 mm S.L., ex Panama, BMNH. 1938.5. 15.305-7. 12. Pellona panamensis Steindachner, 1875 = Ilisha furthii (Steindachner, 1875) Pellona panamensis Steindachner, 1875, Sitzb. K. Akad. Wiss. Wien, 70 : 389 (Panama) ; Idem, 1875, Ichthyol. Beitr., No. i : 15 (repeat). TYPE MATERIAL. LECTOTYPE, a fish of 280 mm S.L. (365 mm tot.l.), ex Pana- ma, NMV.I887 (labeUed 1874-1/909- pt. a.). PARALECTOTYPE, a fish of 265 mm S.L., ex Panama, NMV. 1890 (labelled 1874-1/909- pt.). There is a third, non-typical, specimen of 258 mm S.L., ex Tumbez, coll. Stokm., NMV.I886. DESCRIPTION. Two fishes, 280 and 265 mm S.L., LECTOTYPE and PARA- LECTOTYPE, ex Panama, in fair condition, caudal tips broken, many scales lost, occipital region dissected in paralectotype, NMV.i887 and 1890. (Figures for lecto- type cited first). Br.St. 5 (6), D iii 13 (14), P i 14 (14), V i 6 (6), A iii 49 (47), g.r. 11+24 (11+24), scutes 24+13 (23+13). In percentages of standard length : body depth 32-6 (32-0), body width 8-7 (9-1), head length 26-0 (26-1) ; snout length 7-4 (6-9), eye diameter 7-4 (7-6), length of upper jaw 13-5 (13-1), length of lower jaw 14-2 (13-9) ; pectoral fin length 17-8 (16-7), pelvic fin length 5-9 (6-0), length of anal base 34-4(41-3) ; pre-dorsal distance 49-0 (48-7), pre-pelvic distance 46-0 (45-7), pre-anal distance 62-7 (63-7). These two specimens differ from the type of Ilisha furthii, a smaller fish (201 mm S.L.), only in the more slender body (32-0-32-6 ; cf. 36-4), less convex belly profile, STEINDACHNER'S CLUPEOID FISHES 25 slightly shorter pectoral fins (167-17-8 ; cf. 197) which do not reach the pelvic fin base, and shorter anal fin base (34-4-41-3 ; cf. 45-0). The latter is surprising, especi- ally since the lectotype of P. furthii has fewer anal finrays, but there seems to be no justification for separating Steindachner's P. panamensis, as Norman (1923^ also concluded. PELLONA Valenciennes, 1847 Pellona Valenciennes, 1847, Hist. Nat. Poiss., 20 : 300 (Type : Pellona orbignyana Val., desig- nated by Gill, 1861, Proc. A cad. nat. Sci. Philad. : 38). Neosteus Norman, 1923, Ann. Mag. nat. Hist. (9) 11 117 (Type : Pellona ditchela Valenciennes by subsequent designation of Norman, Zoo/. Rec. Pisces for 1923 : 25). As in the case of Ilisha, the species of Pellona are very much in need of revision. The work of Hildebrand (1964) and examination of the status of the two Valenciennes species, P. castelnaeana and P. flavipinnis (Whitehead, I967a : 106-110), can be sum- marized in the following key. I INDO-PACIFIC AREA ; scutes 18-19 + 8 ; anal iii-iv 32-24 ; gillrakers on lower part of ist arch 23-25 i. p. ditchela Val., 1847 II NEW WORLD A. Post-pelvic scutes 10-14 ; anal "i 32~37 ....... 1. Gillrakers on lower part of ist arch 12-14 (Amazon system) . 2. P. castelnaeana Val., 1847 2. Gillrakers on lower part of ist arch 23-31 (as above but also from Surinam and Parana) 3. P. flavipinnis Val., 1837 B. Post-pelvic scutes 5-7 ; g.r. 23-25 1. Anal iii 33-39 (Atlantic coasts from Panama to southern Brazil) 4. P. harroweri (Fowler, 1917) 2. Anal iii 42 (single specimen from Newport, Rhode Island) 5. P. narragansetae (Fowler, 1911) Pellona ditchela, the single Old World species, not only is inseparable from the New World species on any feature that could be regarded as of generic or subgeneric importance, but is so close to Pellona harroweri of Costa Rican and Brazilian waters that separation of the two is very difficult. Since the only pristigasterine linking these two regions is a species of Ilisha (I. africana of West Africa), it might be assumed that Pellona ditchela and P. harroweri are isolated relicts from a formerly much wider distribution of Pellona. Speciation in the Indo-Pacific may have been held back by competition from the more numerous species of Ilisha, whereas in the New World it is Pellona that appears to have speciated at the expense of Ilisha. The possibility cannot be ruled out that the combination of scute, anal finray and gillraker counts that characterizes P. ditchela and P. harroweri and distinguishes them from all other species may have been arrived at independently in the two species. The use of gillraker numbers to separate P. castelnaeana (12-14) from P. flavipinnis (23-31) seems justified by the discontinuity shown in Table 3 and Figure 2. In an earlier paper (Whitehead, ig67a : no) the low gillraker count in the smaller syntype of P. castelnaeana appeared to be the only exception to a general trend of reduction in number in larger fishes. Inclusion of the low gillraker counts found in three 26 P. J. P. WHITEHEAD specimens (220-275 mm S.L.) of P. altamazonica by Hildebrand (1964 : 418) and the high counts found in large fishes from the Guianas, however, suggests that there are two species, each with fairly constant numbers of gillrakers. A " slight " difference was found in the length of the pectoral axillary scale relative to fin length between the two species (Whitehead, ig67a : no), but the relationship appears to be allo- inetric and overlap occurs. 35 n fip 'o o 25- 20- 5- « o oeo e GO oo - o ® Argentina O Brit., Dutch Guiana O Amazon • SYNTYPES of P. castelneana (Amazon] iii! Rosario (5 fishes) Hildebrand's I. altamazonica 1 r~ ~1~~ ~I~ ~^~ ™« 100 200 300 400 500 S.L. in mm. FIG. 2. Gillraker counts (lower arm, ist arch) in specimens of Pellona flavipinnis (upper series) and P. castelnaeana (lower series). See Table 3, p. 44. As Hildebrand (1964) supposed, P. narragansetae (known only from the holotype from Newport, Rhode Island, i.e. well outside the recorded range of the genus) was probably a stray and its separation from P. harrow eri is perhaps not justified. The following specimens have been labelled types of what appear to be MS. names never published by Steindachner. " Pellona staudingeri " = Pellona flavipinnis (Valenciennes, 1837) A fish of ii8-2 mm S.L., ex Iquitos (Peruvian Amazon), NMV.HI2 (labelled 1884 I 300 a TYPE?). A fish of 105-0 mm S.L., same locality, NMV.i893 (labelled I 1884 300). STEINDACHNER'S CLUPEOID FISHES 27 The specimens have iii 34 -iv 33 anal rays, 15 + 28 and 16 + 29 gillrakers, and 20+ 12 scutes. They are clearly Pellona flavipinnis, the high gillraker count distin- guishing them from P. castelnaeana according to the key given here. " Pellona macrolepis " = Pellona flavipinnis (Valenciennes, 1837) A fish of 340 mm S.L., ex Teffe (middle Amazon), NMV.noi (labelled 1874 — I, TYPE?). This specimen has iii 37 anal rays, 14+26 gillrakers and 23+13 scutes. This high gillraker number reinforces the impression that there is no reduction in large fishes (Figure 2). ODONTOGNATHUS Lacepede, 1800 Odontognathus Lacepede, 1800, Hist. Nat, Poiss., 2 : 220 (Type : Odontognathus mucronatus Lacepede, 1800). Gnathobolus Schneider, 1801, Syst. Ichthyol. Blochii, 2 : 556 (Type : O. mucronatus Lacepede). Odontognathus is separated from the rather similar Opisthopterus Gill by its much longer maxilla (to gill opening or beyond ; cf . to vertical from eye centre in Opisthop- terus). Berry (1964 : 729) showed that the maxilla exhibits positive allometry with standard length, and that in Odontognathus of 55-75 mm the elongation has already begun ; by 100-110 mm the tip of the maxilla has reached the opercular margin. Both genera lack pelvic fins but are otherwise similar to Ilisha. Three species of Odontognathus are currently recognized and can be distinguished by the following key. 1 Scutes interrupted in area below pelvic fin base ; scutes with a single spine, outer margin not serrated ; anal rays 71-82 (Guianas to Trinidad) i. O. mucronatus Lacepede, 1800 2 Scutes in continuous series ; outer margin of posterior scutes with 2-8 serrations a. Anal rays 58-62 ; scutes 25-27 ; dorsal origin over about i6th anal ray (Trinidad to Costa Rica). . . . . . . 2. O. compressus Hildebrand, 1923 b. Anal rays 65-68 ; scutes 29-30 ; dorsal origin over about 25th anal ray (Pacific coast of Panama) . . . . 3. O. panamensis (Steindachner, 1876) 13. Pristigaster (Odontognathus) panamensis Steindachner, 1876 = Odontognathus panamensis (Steindachner, 1876) Pristigaster (Odontognathus) panamensis Steindachner, 1876, Sitzb. K. Akad. Wiss. Wien., 74 : 72 (Panama) ; Idem, 1876, Ichthyol. Beitr., No. 5 : 24 (repeat). TYPE MATERIAL. HOLOTYPE, a fish of 179-0 mm S.L., ex Panama, NMV.4626 (labelled 1874 I 2198 Steind. don.). DESCRIPTION. A fish, 179-0 mm S.L. (204 mm tot.l., estimated), HOLOTYPE, ex Panama, in fair condition but tips of dorsal, anal and caudal fins damaged, scales mostly absent, NMV.4626. 28 P. J. P. WHITEHEAD Br. St. 4 (both), D i n, P i n, V (nil), A i 65, g.r. 8+20, scutes 30. In percentages of standard length : body depth 25-3, body width 5-5, head length 17-3 ; snout length 4-6, eye diameter 4-8, upper jaw length 13-8, lower jaw length 8-0 (height 4-9) ; pectoral fin length 19-5, pelvic fin (absent), length of anal base 51-1 ; pre-dorsal distance 73-8, pre-anal distance 46-8. Body strongly compressed, its width 5 times in its depth, dorsal profile " humped " at nape (? partly distorted) and concave over eye ; belly strongly keeled, scutes begin- ning on hind part of isthmus, prominent throughout, margins of nth, I2th, i7th-3Oth scutes with 2-8 serrations (Figure 3b). FIG. 3. Pristigaster (Odontognathus) panamensis HOLOTYPE, 179 mm S.L. NMV-4626 ( = Odontognathus panamensis). a. Head and anterior half of body. b. Detail of serrated scutes behind pectoral fin tips. Eye about equal to snout length, 3^ times in head length. Upper jaw reaching to just beyond gill opening ; median £ of upper jaw edentulous, a single series of 10-12 fine, conical teeth on either side of this ; no hypo-maxilla ; lower edge of maxilla with a series of minute teeth reaching to posterior tip of bone ; two supra-maxillae present, the ist (anterior) elongate kidney-shaped, 4-1 mm in length, the 2nd (posterior) with diamond-shaped expanded posterior part, 6-0 mm long and 3-9 mm deep, its posterior tip lying below vertical from eye centre ; maxilla depth rapidly decreasing behind tip of 2nd supra-maxilla, thereafter forming a slender blade (Figure 3a). Lower jaw slightly projecting, upper border rising rapidly, its height STEINDACHNER'S CLUPEOID FISHES 29 reaching about | of jaw length in first third of jaw ; a single series of about 5-6 small conical teeth on either side of symphysis. Operculum with lower border rising at about 45° ; sub-operculum twice as long as deep. Gill opening entirely covered by opercular series. No cleithral lobe. Snout with strong, blade-like mid-dorsal ridge dividing before eyes and extending to hind end of skull ; two smaller ridges on either side, enclosing a canal ; a median ridge at hind end of skull and two faintly striated fronto-parietal triangular areas. Pseudobranch present, exposed, with 10 filaments, its length 5-5 mm (£ eye dia- meter). Gillrakers slender, the longest 4-4 mm (f eye diameter) ; gill filaments shorter, the longest 3-3 mm. Two stubby gillrakers present on posterior face of 3rd epibranchial (none on 2nd arch). Granular teeth present on tongue, palatines, ecto- and endo-pterygoids, the toothed area becoming ridged posteriorly in the latter. Dorsal fin origin above 25th branched anal ray, twice as near to caudal base as to posterior margin of pre-operculum ; dorsal rays damaged at tips, but apparently only a single, rather broad unbranched ray ; dorsal base short, f of eye diameter. Pectoral fin broad, its tip failing to reach anal origin by 2 eye diameters ; no axillary scale present. Pelvic fins absent. Anal fin base £ standard length, anal origin nearer to snout than to caudal base by i£ eye diameters. Scales with slightly eroded posterior border and a few faint radiating striae ; un- exposed portion without the main, uninterrupted vertical striation characteristic of clupeoid scales, but with 3-4 faint radiating striae not reaching centre of scale. Colour : general light brown with two faint longitudinal silver lines, the first midlateral and the second a little above it ; cheek, operculum and belly silvery. IDENTIFICATION. Odontognathus panamensis is very similar to 0. compressus Hildebrand. Hildebrand (1923 : 194) found the dorsal origin slightly further forward in 0. compressus, the anal fin base shorter and with fewer rays (58-62 ; cf. 65-68 in 0. panamensis), and fewer ventral scutes (25-27 ; cf. 29). These are fairly small differences and a large sample of 0. panamensis might bridge the discontinuity in finray and scute numbers. Both species have serrated scutes, which separates them from 0. mucronatus, the latter further distinguished by a short non-scuted area below the pectoral fin base. Odontognathus compressus may yet prove to be merely an Atlantic subspecies of the Pacific 0. panamensis. Family ENGRAULIDAE ANCHOVIELLA Fowler, 1911 Anchoviella Fowler, 1911, Proc. Acad. nat. Sci. Philad., 63 : 211 (Type : Engraulis perfasciatus Poey ) . Hildebrand (1943 : 108) redefined this genus to include species with a short and posteriorly rounded maxilla (Fowler had stressed the low number of gillrakers) ; Hildebrand recognized the subgenus Amplova Jordan & Scale for three species with exceptionally short maxillae, A. balboae (Jordan & Scale), A.jamesi (Jordan & Scale) and A . brasiliensis Hildebrand. Concentrating on maxilla length, Hildebrand failed 3o P. J. P. WHITEHEAD to recognize the nominal species Stolephorus eury stole Meek & Swain and Anchoviella estanquae Hildebrand as members of Engraulis (Whitehead, 1964^. Anchoviella pallida (Starks) belongs to another genus (see under Anchovia, p. 38), leaving 16 species of Anchoviella, distinguished mainly by rather small meristic and morpho- metric differences. In the majority of these species the number of gillrakers on the lower arm of the first arch is in the range 15-28 ; A. balboae, with 29-35, stands out- side this range but examination of paratypical specimens of Anchovia brevirostra ( = A . balboae) confirms this count and shows that this species is otherwise a fairly typical member of the Amplova group. 14. Engraulis vaillanti Steindachner, 1908 = Anchoviella vaillanti (Steindachner, 1908) Engraulis vaillanti Steindachner, 1908, Anz. Akad. Wiss. Wien, 45 : 193 (Joazeiro and Barra on Rio Sao Francisco, Rio Grande do Norte, and Rio Preta, Brazil). TYPE MATERIAL. a. LECTOTYPE, a fish of 62-7 mm S.L., ex Fazenda Ingaziera, labelled 1903 23/4b, NMV.I93I. b. PARALECTOTYPES, 15 fishes, 39-68 mm S.L., from the same jar. c. PARALECTOTYPES, 16 fishes, 43-0-59-0 mm S.L., ex Lagao Viana, labelled 1903, MNV.I928. d. PARALECTOTYPES, 6 fishes, 42-0-51-5 mm S.L., ex Lagao do Porto, labelled 1903 (£)a, NMV.I929. e. PARALECTOTYPES, 3 fishes, 42-9-45-0 mm S.L., ex Rio Preto, labelled I903a, NMV.i-930. f. PARALECTOTYPES, 4 fishes, 39-2-42-0 mm S.L., ex Lagao Viana and Lagao do Porto, labelled 1903 31/3 £, NMV.I932. g. PARALECTOTYPES, 2 fishes, 39-2-40-6 mm S.L., ex Lagao do Porto, labelled I9O3C, NMV.I938. h. PARALECTOTYPES, 4 fishes, juveniles of 25-0-37-2 mm S.L., ex Barinha, labelled 1903 17/3 b, NMV.I936. DESCRIPTION. A fish, 62-7 mm S.L. (75-3 mm tot.l.), LECTOTYPE, ex Fazenda Ingaziera, in fair condition but scales mostly lost, lower caudal lobe broken, NMV. Br.St. 12, D in 10, P i 12, V i 6 (both), A iii 20, g.r. 14+19. In percentages of standard length : body depth 21-2, body width 9-2, head length 26-3 ; snout length 5-1, eye diameter 7-1, length of upper jaw 20-5, length of lower jaw 16-7 ; pectoral fin length 16-7 (axillary scale 8-7), pelvic fin length n-6 (axillary scale 4-7), length of anal fin base 23-5 ; pre-dorsal distance 54-5, pre-pelvic distance 45-5, pre-anal distance 65-8. Body fairly compressed, its width 2-4 times in its depth, the latter less than head length. Snout moderately prominent, a little less than eye diameter. Width of head above eye a little less than eye diameter. Upper jaw just reaching to mandi- bular articulation but not quite to anterior border of pre-operculum ; tip of maxilla STEINDACHNER'S CLUPEOID FISHES 31 evenly rounded, projecting only slightly beyond tip of 2nd (posterior) supra-maxilla ; the latter spatulate, tapering anteriorly and overlain half-way along its length by a plate-like ist (anterior) supra-maxilla. Fine, close-set teeth on edge of maxillae, pre-maxillae and dentaries and minute teeth on palatines and ecto- and endoptery- goids, but not on vomer. Gillrakers fine, slender, the longest 2 mm long (about £ eye) and i£ times length of corresponding gill filaments ; 9 short rakers on posterior face of 3rd epibranchial. Pseudobranch present, exposed, about £ eye diameter. Isthmus brown (silvery in life?), sterno-hyoideus muscle not quite reaching to posterior border of branchio- stegal membrane, the ventral edge of the urohyal exposed before this. A pair of crescentic posterior frontal fontanelles, together forming a triangle 1-6 mm long and 1 7 mm wide. Dorsal fin origin equidistant between caudal base and posterior pupil border, i.e. nearer to caudal base by i| eye diameters. Pectoral fin tips failing to reach pelvic base by | eye diameter ; axillary scale present, almost £ length of fin. Pelvic fin base $ eye diameter before vertical from dorsal origin, nearer to pectoral base than to anal origin by \ eye diameter ; axillary scale present, f length of fin ; inner rays of fins joined by membrane. Anal fin with low scaly sheath, its origin just behind vertical from last dorsal ray and \ eye diameter nearer to pectoral than to caudal bases. Scales : unexposed portion with well-defined " shoulders " and 7-8 short horizontal striae, the two median striae meeting an irregular vertical striation ; exposed portion of scale with an irregular vertical striation followed by a reticular pattern of striae covering the rest of the posterior part of the scale ; hind border of scale not eroded. In some scales the reticular pattern extends forward and disrupts the (apparently) normal pattern of striae. Colour : body brown with faint suggestion of silvery midlateral stripe ; a peppering of melanophores on snout, along entire dorsal profile, along posterior half of mid- lateral band (becoming heavier posteriorly) and on dorsal fin ; a dark vertical bar at base of caudal. NOTE. Hildebrand (1943) had no specimens of A. vaillanti, but on the basis of Steindachner's description he distinguished the species by its high number of anal finrays, relatively few gillrakers and dorsal origin nearer to snout tip than to caudal base. As far as the lectotype is concerned, the latter is not true. Except for the posteriorly placed anal origin, A . vaillanti is close to A . Upidenstole (Fowler) but it has a more slender body. It also resembles A. hubbsi Hildebrand but has fewer gillrakers and a less prominent silver lateral band. 15. Engraulis nattereri Steindachner, 1879 = Anchoviella nattereri (Steindachner, 1879) Engraulis nattereri Steindachner, 1879, Sitzb. K. Akad. Wiss. Wien, 80 : 174 (Para, Brazil) ; Idem, 1879, Ichthyol. Beitr., No. 8 : 56 (repeat). TYPE MATERIAL. Intensive search failed to produce the type, a specimen of 50 mm. It may have been sent with duplicates to another museum. 32 P. J. P. WHITEHEAD DESCRIPTION, (based on Steindachner's description). Br.St. (n.r.), D 12, P(n.r.), V (n.r.), A 28 or 29, g.r. (n.r.). Snout fairly long, reaching well beyond tip of lower jaw, 4-0 in head ; eye larger than snout, 3-6 in head. Maxilla tip nearly square, not quite reaching to mandi- bular articulation. Gillrakers moderately long, longest equal to eye diameter. Dorsal fin origin slightly nearer to base of caudal than to tip of snout. Pectoral fin a little longer than postorbital distance, its tip reaching half-way along pelvic fin. Anal fin origin below middle of dorsal base. Colour : silver midlateral band indistinct. NOTE. Maxilla shape, number of anal finrays and position of anal origin all strongly suggest a species of Anchovietta. Hildebrand (1943 : 133), who had no specimens and relied solely on Steindachner's description, kept this species distinct. The rather long pectoral fins suggest A. pallida (Starks), but with no record of gill- raker number it seems best to follow Hildebrand for the moment. ANCHOA Jordan & Evermann, 1927 Anchoa Jordan & Evermann, 1927, Proc. Calif. Acad. Sci., (4) 16 (15) : 501 (Type : Engraulis compressus Girard). This is the largest engraulid genus, with over thirty species recognized by Hilde- brand (1943). The genus was originally distinguished from the very similar Ancho- viella by the possession of more anal rays and gillrakers. Tables of anal ray and gillraker numbers given by Hildebrand (loc. cit.} show considerable overlap, however, and Hildebrand redefined the genus on the basis of the length of the maxilla : Anchoa : tip of maxilla pointed, projecting well beyond tip of 2nd supra-maxilla and reaching beyond mandibular articulation, almost to gill opening. Anchovietta : tip of maxilla truncate or bluntly rounded, projecting only slightly beyond tip of 2nd supra-maxilla and not reaching beyond mandibular articulation. Three principal features distinguish both Anchoa and Anchoviella from Engraulis (Whitehead, ig6^b ; summarized in Berry, 1964). 1. Posterior frontal fontanelles present (occluded in adult Engraulis). 2. Anal origin below or only just behind vertical from last dorsal ray (up to one eye diameter behind in Engraulis). 3. Pseudobranch short, equal to or less than eye diameter, not reaching to hyo- mandibular facet nor onto inner face of operculum, 15-25 pseudobranchial filaments (greater than eye, etc., 25-40 filaments in Engraulis). Using these criteria, Anchoviella eury stole (Swain & Meek) and A. estanquae Hildebrand were recognized as members of Engraulis (Whitehead, I9&4b : 882). The redescription of the type of Anchoa nasus given below (p. 34) poses a further problem. In the three characters listed above, A. nasus approaches Engraulis while still maintaining the diagnostic feature of Anchoa, the long maxilla. The anal fin origin is not so far back as in Engraulis, the posterior frontal fontanelles appear to be excluded rather later in ontogeny and the pseudobranch is slightly shorter than STEINDACHNER'S CLUPEOID FISHES 33 in Engraulis, but A . nasus is as close to Engraulis in these respects as it is to typical members of Anchoa. It has been noted (Whitehead, I96ya : 127) that three further species of Anchoa share these resemblances to Engraulis, viz. A. lyolepis (Evermann & Marsh), A. argentivittata (Regan) and A. duodecim (Cope) (B.M. specimens, includ- ing the type of A. argentivittata}. Full revision of the anchovy genera may well support recognition of a separate genus or subgenus for these four species. 16. Engraulis januariusSteindaichner, 1879 = Anchoa januaria (Steindachner, 1879) Engraulis januarius Steindachner, 1879, Stizb. K. Akad. Wiss. Wien, 80 : 176 (ex harbour of Rio de Janeiro, Brazil) ; Idem, 1879, Ichthyol. Beitr., No. 8 : 58 (repeat). TYPE MATERIAL. LECTOTYPE, a fish of 51-4 mm S.L., ex Rio de Janeiro har- bour, labelled " 1874 I 1566 ", NMV.279O. PARALECTOTYPE, a fish of 54 mm S.L., from same jar. DESCRIPTION. A fish, 51-4 mm S.L., LECTOTYPE, ex Rio de Janeiro harbour, in good condition, NMV.279O. Br.St. n, D iii 13, P i 12, V i 6, A iii 20, g.r. 21+27, scales ca 37 in lateral series. In percentages of standard length : body depth 21-2, body width 7-8, head length 23-2 ; snout length 4-9, eye diameter 7-0, length of upper jaw 21-0, length of lower jaw 16-2 ; pectoral fin length 13-2, pelvic fin length 7-2, length of anal base 21-6 ; pre-dorsal distance 56-5, pre-pelvic distance 45-9, pre-anal distance 64-0. Body compressed, its width 3 times in depth, the latter slightly less than head length. Snout a little shorter than eye diameter. Upper jaw reaching back beyond articulation of lower jaw and to vertical midway across pre-operculum ; lower edge of maxilla straight posteriorly, tip rounded, upper edge rounded to meet posterior tip of 2nd supra-maxilla ; the latter spatulate, tapering anteriorly ; anterior (ist) supra- maxilla slender, plate-like ; the maxilla projects 1-7 mm beyond the tip of the 2nd supra-maxilla. Teeth on pre-maxillae, maxillae and dentaries close-set, fine, very short ; fine teeth on palatines and ecto- and endo-pterygoids but not on vomer. Gillrakers fine, slender, twice length of corresponding gill filaments and f eye dia- meter ; 8 short rakers on posterior face of 3rd epibranchial. Pseudobranch present, partly invested in adipose tissue, with about ten short filaments, the length of the pseudobranch ^ eye diameter. Isthmus silvery, sterno-hyoideus muscle extending forward almost to hind margin of branchiostegal membrane, the ventral edge of the urohyal exposed before this. Width of head over eye centre equal to eye diameter. A pair of triangular posterior frontal fontanelles, together 1-5 mm long and 1-4 mm at widest (posterior) point. Dorsal fin origin equidistant between caudal base and posterior pupil border, i.e. nearer to caudal base than to snout tip by i^ eye diameters ; proximal half of fin invested in scaly sheath. Pectoral fin tips failing to reach pelvic base by £ eye diameter ; axillary scale present, -£$ length of fin. Pelvic fins small ; pelvic base 1 1 eye diameters before vertical from dorsal origin and equidistant between pectoral base and anal origin ; large axillary scale present (missing on right side), equal to fin 34 P. J- P. WHITEHEAD length. Anal fin origin below vertical from base of 8th branched dorsal ray and slightly nearer to caudal than to pectoral bases ; proximal half of fin invested in scaly sheath. Scales : thin, apparently not caducous ; exposed portion with three complete irregular vertical striae, posterior edge slightly eroded ; unexposed portion with two short, incomplete vertical striae and two or three short radial striae from anterior edge of scale. The circulae on the unexposed portion of scale more widely spaced than those on the exposed portion ; the posterior ^ of scale apparently without circulae. Colour : body brown, with faint silvery midlateral stripe and silvery belly ; dark chromatophores forming diagonal line on upper part of caudal peduncle and a vertical bar below this. NOTE. Hildebrand (1943) listed 36 species of Anchoa, for which he provided a complex and not altogether satisfactory key. Several species resemble Anchoa januaria, notably A. mitchilli, A. parva, A. hepsetus, A. pectoralis and A. tricolor (lectotype of the first fully described in Whitehead, ig67a : 127). Assessment of the importance of the small and mainly meristic differences separating these species must await full revision based on more material. 17. Engraulis nasus Kner & Steindachner, 1866 = Anchoa nasus (Kner & Steindachner, 1866) (Plate 3a) Engraulis nasus Kner & Steindachner, 1866, Sitzb. K. Akad. Wiss. Wien, 54 : 388, pi. 2 (17) (Chincha I., Peru). Stolephorus cultratus Gilbert, 1892, Proc. U.S. natl. Mus., 14 : 544 (Santa Margarita I., off Lower California) . SYNONYMY. Hildebrand (1943 : 70), who noted that the type of Stolephorus cultratus cannot now be found, allied that species with Anchoa ischana (Jordan & Gilbert) and A. naso. From Gilbert's description, however, 5. cultratus is even closer to Anchoa nasus, differing only in minor respects (anal 20 ; cf. 21-27 ; maxilla " nearly to gill opening " ; cf. " not to margin of opercle "). Gilbert's species is perhaps a northern representative of Anchoa nasus. TYPE MATERIAL. HOLOTYPE, a fish of 107-8 mm S.L., ex Chincha I., Peru, labelled Steindachner 1866 67, NMV.2837. DESCRIPTION. A fish, 107-8 mm S.L. (132-6 mm tot.l.) HOLOTYPE, ex Chincha I., Peru, in good condition but scales mostly lost, NMV.2837. Br.St. 13, D iii 13, P i 13, V i 6, A iii 20, g.r. 23+27. In percentages of standard length : body depth 23-2, body width 8-9, head length 30-0 ; snout length 6-0, eye diameter 7-4, length of upper jaw 27-5, length of lower jaw 19-8 ; pectoral fin length 16-0 (axillary scale 9-5), pelvic fin length 10-2, length of anal base 22-6 (height of fin 12-2) ; pre-dorsal distance 56-2, pre-pelvic distance 45-8, pre- anal distance 69-0. Body compressed, its width 2-6 times in its depth, the latter f of head length. STEINDACHNER'S CLUPEOID FISHES 35 Snout prominent, a little less than eye diameter. Upper jaw reaching back almost to hind margin of pre-operculum ; tip of maxilla pointed ; two supra-maxillae, the 2nd (posterior) spatulate, reaching to anterior margin of pre-operculum, the ist (anterior) slender, plate-like ; maxilla projects 3-85 mm beyond tip of 2nd supra- maxilla. Anterior tip of lower jaw below vertical from anterior eye border. Teeth on entire lower edges of maxillae and on dentaries, fine and fairly close-set ; teeth on pre-maxillae even finer, granular teeth on palatines, ecto- and endo-pterygoids but not on vomer. Gillrakers fine, slender, i^ times length of corresponding gill filaments and ^ eye diameter ; 9 short, stubby rakers on posterior face of 3rd epibranchial. Pseudo- branch present, fully exposed, its length i| eye diameters, about 30 filaments, the last two reaching onto the inner face of the operculum. Isthmus silvery, sterno- hyoideus muscle extending forward to posterior border of branchiostegal membrane, exposed portion of urohyal in front of this. Width of head over eye centre a little greater than eye diameter. Posterior frontal fontanelles occluded, the posterior tips of the frontals meeting in the midline and dividing only after reaching the supra- occipital ; frontal tips rounded posteriorly. Dorsal fin origin equidistant between caudal base and posterior pupil border, i.e. nearer to caudal base than to snout tip by if eye diameters ; the few remaining scales suggest a scaly sheath to the base of the fin. Pectoral fin tips reaching £ eye diameter beyond pelvic base ; axillary scale present, $ length of fin. Pelvic fin base i£ eye diameters before vertical from dorsal origin, \ eye diameter nearer to pectoral base than to anal origin ; inner rays joined to each other by a membrane, overlain by two elongate scales ; no axillary scale found. Anal origin nearer to caudal than to pectoral base by \ eye diameter ; first unbranched anal ray a fraction behind vertical from base of last dorsal ray. Base of caudal with elongate scales reaching half-way along fin. Scales : thin, many missing from anterior part of body but remainder firmly fixed ; anterior part of scale with distinct 'shoulders ' and 7 horizontal striae whose ends bend towards the centre of the scale ; exposed portion of scale with (usually) seven vertical pairs of striae, the first two short, the third bent posteriorly half way along their lengths, the fourth short, the fifth meeting in the centre of the scale and the remaining two curved and either short or continuous across the scale (in many scales, possibly regenerated scales, the striae are reticulate, more or less disrupting the apparently normal pattern of striation). Colour : upper } of body light brown, remainder silvery, no sign of silver lateral stripe. NOTE. The descriptions given by Hildebrand (1943) are insufficient to judge whether the resemblances of A. nasus to A. ischana and A. naso are superficial or whether they include the three features which A. nasus shares with A. lyoUpis, A . argentivittata and A . duodecim, and which serve to distinguish these four species from all other members of Anchoa (see under genus, p. 32). In the original description, the specimen was said to have a strongly convex dorsal profile and a nearly straight ventral profile (see Plate 3a). Hildebrand (1943 : 104) correctly assumed this to be a preservation artifact. 36 P. J. P. WHITEHEAD 18. Engraulis peruanus Steindachner, 1879 = Anchoa nasus (Kner & Steindachner, 1866) Engraulis peruanus Steindachner, 1879, Sitzb. K. Ahad. Wiss. Wien, 80 (ex Callao, Peru) ; Idem, 1879, Ichthyol. Beitr., No. 8 : 60 (repeat). TYPE MATERIAL a. LECTOTYPE, a fish of 100-0 mm S.L., ex Callao, labelled 1874 I 1215 (pt.a) Steind. don., NMV.IQ65. b. PARALECTOTYPES, 3 fishes, 91-0-102-4 mm S.L., ex Callao, from same jar. c. PARALECTOTYPE, i fish, 94-8 mm S.L., ex Callao, labelled as above but (pt.d), NMV.I966. d. PARALECTOTYPES, 5 fishes, 79-3-97-6 mm S.L., ex Paraca Bay, as above, NMV.I964. e. PARALECTOTYPES, 2 fishes, 86-4-89-4 mm S.L., ex Callao, as above but (pt.), NMV.I967. f. PARALECTOTYPES, 2 fishes, 85-3-87-4 mm S.L., ex Callao, as above but (pt.c), NMV.I965. DESCRIPTION. A fish, 100 mm S.L. (122-0 mm tot.l.), LECTOTYPE, ex Callao, in good condition but some scales lost, NMV. 1965. Br.St. 13, D iii 13, P i 13, V i 6, A iii 23, g.r. 23+27. In percentages of standard length : body depth 24-6, body width 8-3, head length 29-4 (width above eye centre 7-0) ; snout length 5-2, eye diameter 7-2, length of upper jaw 27-3, length of lower jaw 20-9 ; pectoral fin length 16-6 (axillary scale 9-8), pelvic fin length 10-0 (axillary scale 8-5), length of anal base 23-2 ; pre-dorsal dis- tance 54-5, pre-pelvic distance 46-8, pre-anal distance 66-1. In the remaining proportional and other features this specimen closely resembles the holotype of Anchoa nasus except that the anal fin origin is slightly further for- ward (below nth branched dorsal ray ; cf. just behind last (i3th) branched dorsal ray). NOTE. Hildebrand (1943 : 102) placed Engraulis peruanus in the synonymy of Anchoa nasus on the basis of nine Steindachner specimens from Callao in the Museum of Comparative Zoology at Harvard. The description given here reinforces Hilde- brand's view. 19. Engraulis panamensis Steindachner, 1875 = Anchoa panamensis (Steindachner, 1875) Engraulis panamensis Steindachner, 1875, Sitzb. K. Akad. Wiss. Wien, 72 : 589 (ex Panama) ; Idem, 1875, Ichthyol. Beitr., No. 4 : 39 (repeat). TYPE MATERIAL a. LECTOTYPE, a fish of 113-2 mm S.L., ex Panama, labelled 1874 I 1149 (pt.) Steind., NMV.I97O. b. PARALECTOTYPES, 2 fishes 112-7-125-0 mm S.L., ex Panama, from the same jar. STEINDACHNER'S CLUPEOID FISHES 37 c. PARALECTOTYPES, 2 fishes, 113-6-113-8 mm S.L., ex Panama, as above, NMV.I969. d. PARALECTOTYPES, 2 fishes, 109-2-114-9 mm S.L., ex Panama, as above, NMV.I972. e. PARALECTOTYPES, 2 fishes, 91-2-97-5 mm S.L., ex Panama, as above, NMV.I97I. DESCRIPTION. A fish, 113-2 mm S.L. (139 mm tot.l., estimated, caudal tips broken), LECTOTYPE, ex Panama, in fair condition, NMV.I970. Br.St.i2, D iii n, P i 13, V i 6, A iii 31, g.r. 18+22. In percentages of standard length : body depth 27-4 (width 6-0), head length 22-8 (width over eye centre 6-1) ; snout length 3-5, eye diameter 6-6, length of upper jaw 20-7, length of lower jaw 15-8 ; pectoral fin length 20-2 (axillary scale 5-7), pelvic fin length 9-2, length of anal base 35-2 ; pre-dorsal distance 56-8, pre-pelvic distance 41-2, pre-anal distance 56-3 ; caudal peduncle, length 12-4, depth 10-2. Body strongly compressed, its width 4-5 times in its depth, the latter greater than head length. Head rather deep and dorsally (snout to supra-occipital) rather short. Snout not strongly prominent, almost half eye diameter. Upper jaw reaching almost to gill opening (right maxilla tip bent downward, left maxilla tip possibly lacking final mm) ; tip of maxilla apparently sharply pointed, projecting well beyond 2nd (posterior) supra-maxilla, the latter spatulate and tapering anter- iorly ; ist (anterior) supra-maxilla plate-like. Symphysis of lower jaw slightly before vertical from anterior eye border. Teeth along entire lower edges of maxillae and on dentaries, fine and close set ; teeth on pre-maxillae very small ; granular teeth on palatines, ecto- and endo-pterygoids and a patch of 4 small teeth on either side of the vomer ; a line of granular teeth along upper edge of ceratohyal. Gillrakers fine, slender, f eye diameter and twice length of corresponding gill filaments ; 6 short rakers on posterior face of 3rd epibranchial. Pseudobranch pres- ent, exposed, £ eye diameter. Isthmus silvery, sterno-hyoideus muscle extending forward just to posterior margin of branchiostegal membrane, urohyal exposed before this. Posterior frontal fontanelles exposed, crescentic, separated anteriorly by a wedge, the fontanelles 1-8 mm long and together 2-2 mm wide. Dorsal fin origin almost 2 eye diameters nearer to caudal base than to snout tip ; a low scaly sheath along base. Pectoral fin tips reaching beyond pelvic base by f eye diameter ; axillary scale present, short, about £ length of fin. Pelvic fin base equidistant between pectoral base and anal origin ; axillary scale Plost ; inner rays of fin joined by membrane to body and perhaps originally to each other. Anal origin equidistant between caudal base and posterior border of pupil and directly below vertical from dorsal origin ; low scaly sheath present. Caudal peduncle almost as deep as long. Scales : oval, deeper than wide, not firmly fixed, many missing ; anterior part of scale with ill-defined radiating striae and a single irregular vertical striation traversing scale ; exposed portion with 2-3 pairs of short vertical striae followed by numerous fine vertical striae increasingly connected to each other posteriorly to form a network ; posterior margin of scale apparently eroded. In many instances this pattern is dis- rupted by extensive reticulation of the striae. 38 P. J. P. WHITEHEAD Colour : body light brown except for silvery lateral stripe a little wider than pupil diameter ; opercular series silvery. NOTE. This species is well defined in the key and description given by Hilde- brand (1943) except that in his specimens the silvery lateral band on the body was " nowhere much broader than pupil. " The possession of vomerine teeth is not common in South American anchovies but occurs in some Indo-Pacific engraulids (e.g. Thryssa). The presence of denticula- tions along the upper edge of the ceratohyal, however, appears to be very rare in clupeoid fishes ; it is also found in large Pterengraulis atherinoides (135-190 mm S.L. — discrete tooth plates) and in Anchoa spinifer (over 70 mm S.L. — finely granular edge) ; in Sardinofis there are numerous short plate-like but soft gillrakers. ANCHOVIA Jordan & Evermann, 1896 Anchovia Jordan & Evermann, 1896, Bull. U.S. nail. Mus., No. 47 (i) : 449 (Type : Engraulis macrolepidotus Kner & Steindachner) . This genus includes fishes which bear a strong resemblance to Cetengraulis (deep and compressed body, reduced jaw dentition and numerous close-set gillrakers which increase in number with size of fish). Members of Cetengraulis are clearly distin- guished, however, by the broadly united branchiostegal membranes and the rela- tively long branchiostegal rays (about £ head length ; cf. £-£ in other engraulid genera). A further reason for considering Anchovia close to Cetengraulis is that these two genera share a peculiarity that appears to be of some systematic importance in the clupeoid fishes, namely the absence of gillrakers on the posterior face of the 3rd epibranchial. First noticed by Dr. Carl Hubbs (in litt.) in the New World species of Engraulis (E. ringens, E. mordax, E. anchoita and E. juruensis, but not in E. eury stole and E. estanquae, which belong in Engraulis sensu stricto), the absence of these gillrakers is now confirmed in Hildebrandichthys as well as in the Indo-Pacific engraulid genus Coilia. These rakers are also absent in Gilchristella aestuarius and Ehirava malabaricus (subfamily Pellonulinae) and Clupea (Strangomera) bentincki (subfamily Clupeinae). The genus Anchovia contains five species in which the maxilla tapers to a point and projects markedly beyond the 2nd supra-maxilla, namely A. macrolepidota (Kner & Steind.), A. magdalenae Hildebrand, A. rastralis (Gilbert & Pierson), A. clupeoides (Swainson) and A . nigra Schultz. Three further species closely resemble the above (posterior gillrakers of 3rd epibranchial absent, etc.), but have a blunt maxilla not reaching beyond the mandibular articulation and barely projecting beyond the tip of the 2nd supra-maxilla, namely A. surinamensis (Bleeker), A. pallida (Starks) and A. potiana Schultz & Menezes. Since maxilla shape is the principal distinction between the genera Anchoa and Anchoviella, revisionary work may well justify splitting the genus Anchovia. STEINDACHNER'S CLUPEOID FISHES 39 20. Engraulis macrolepidotus Kner & Steindachner, 1865 = Anchovia macrolepidota (Kner & Steindachner, 1865) (Plate 3b) Engraulis macrolepidotus Kner & Steindachner, 1865, Abh. K. Bayer Akad. Wiss., 10 : 21, pi. 3 (2) (RioBayano, Panama) ; Steindachner, 1876, Sitzb. K. Akad. Wiss. Wien, 72 ^87 (Panama). TYPE MATERIAL a. LECTOTYPE, a fish of 1047 mm S.L., ex Panama, labelled 1876 II/iP, NMV.28o8. b. PARALECTOTYPE, a fish of 90-3 mm S.L., ex Panama, from the same jar. c. PARALECTOTYPE, a fish of 136-6 mm S.L., ex Acapulco, Mexico, labelled 1874 I 1736 Kn. St., NMV.2807. DESCRIPTION. A fish, 1047 mm S.L. (129-2 mm tot.l.), LECTOTYPE, ex Panama, in fair condition but scales mostly lost, NMV.28o8. Br.St. 14, D iii 12, P i 14, V i 6, A iii 27, g.r. 108. In percentages of standard length : body depth 31-7 (width 7-8), head length 29-7 ; snout length 3-9, eye diameter 6-8, length of upper jaw 26-4, length of lower jaw 20-4 ; pectoral fin length 15-1, pelvic fin length 7-5, length of anal base 28-4 ; pre-dorsal distance 54-2, pre-pelvic distance 45-1, pre-anal distance 59-5. Body compressed, its width 4 times in its depth, the latter a little greater than head length. Snout fairly pointed, a little over half eye diameter ; width of head above eye centre less than eye diameter. Upper jaw reaching a little beyond mandi- bular articulation to posterior margin of pre-operculum ; tip of maxilla pointed, projecting 4-0 mm beyond tip of 2nd (posterior) supra-maxilla ; the latter spatulate, tapering evenly anteriorly and overlain about halfway along its length by the smaller ist (anterior) supra-maxilla. Fine denticulations along edges of maxilla and man- dible becoming fainter anteriorly and not present on pre-maxillae ; fine teeth present along edges of palatines and ecto- and endo-pterygoids, but not on vomer. Gillrakers very fine and slender, the longest (7 mm) slightly exceeding eye dia- meter ; a double series of minute serrae along inner face of each raker ; no gillrakers present on posterior face of 3rd epibranchial. Gill filaments very short, 4^ times in length of gillrakers. Pseudobranch present, exposed, equal to eye diameter, bearing 24 filaments. Isthmus silvery, sterno-hyoideus muscle dividing halfway along to expose the ventral edge of the urohyal, the latter covered by silvery tissue until shortly before the hind margin of the branchiostegal membrane. A pair of exposed posterior frontal fontanelles, 1-4 mm in length, lateral borders forming a sigmoid curve. Opercular series (Figure 4) not covering gill opening ; operculum inclined at an angle of 45° ; sub-operculum almost triangular, the junction of the posterior and ventral margins produced into a distinct point. Dorsal fin origin nearer to snout tip than to caudal base by \ eye diameter ; base of fin not invested in scaly sheath (? scales lost). Pectoral fin tips just reaching to pelvic fin base ; axillary scale present, \ length of fin, bearing a narrow flange along lower edge ; pectoral fins set low on body, below level of sub-operculum. Pelvic fins small ; pelvic base i eye diameter before vertical from dorsal origin and equidis- 4o P. J. P. WHITEHEAD tant between base of pectoral and anal origin ; no axillary scale (? lost) ; final rays of fin joined together in midline by a membrane and also similarly joined to body. Anal fin origin below middle of dorsal base (7th branched dorsal ray) and 2 eye dia- meters closer to pectoral base than to caudal base ; base of fin invested in scaly sheath. FIG. 4. Engraulis macrolepidotus LECTOTYPE, 104-7 mm S.L., NMV.28o8 ( = Anchovia macrolepidota) . Opercular series (right side) showing characteristic shape of sub-oper- culum. Scales : mostly missing ; exposed portion with an apparently random pattern of reticulated striae, unexposed portion with a single continuous curved striation pre- ceded by up to five smaller radiating striae most of which fail to reach the centre of the scale. Colour : upper £ of body brown, remainder silvery ; a dark line across bases of first few upper caudal rays ; very dark brown pigmentation on inner face of branchio- stegal membrane. LYCENGRAULIS Giinther, 1868 Lycengraulis Giinther, 1868, Cat. Fish. Brit. Mus., 7 : 385 and 399 (Type : Engraulis grossidens Agassiz. Hildebrand (1943) listed seven species of Lycengraulis and two further species have since been described, L. limnichthys Schultz and L. simulator De Plaza. In his key (p. 141), Hildebrand separated three species on the basis of their short, partly STEINDACHNER'S CLUPEOID FISHES 41 rudimentary, gillrakers (L. abbotti, L. barbouri and L. schroederi). De Plaza (1962) noted a regression in the gillrakers of L. simulator with increasing size of fish but this did not reach the extreme condition found in the three species listed above. Hilde- brand's specimens were all fairly large (148 and 205-237 mm S.L.), but he reported normal gillrakers in specimens of other species of a comparable size. It can be noted that small gillrakers are present on the posterior face of the third epibranchial in most species of Lycengraulis but are reduced or absent in specimens of L. barbouri at about 140 mm S.L. ; possibly this also occurs in the other species in which the gillrakers regress with age (L. abbotti and L. schroederi). 21. Engraulis poeyi Kner & Steindachner, 1865 = Lycengraulis poeyi (Kner & Steindachner, 1865) (Plate 3c) Engraulis poeyi Kner & Steindachner, 1865, Abh. K. Bayer Akad. Wiss., 10 : 23, pi. 3, fig. 3 (Rio Bayano, Panama). TYPE MATERIAL. Intensive search failed to produce any type material. NOTE. Lycengraulis poeyi was well described by Hildebrand (1943 : 146), who placed it in the group with rather long and numerous gillrakers. It is the only member of the genus reported from the Pacific coasts of Central and South America ; it is well distinguished from the related L. grossidens and L. olidus of the Atlantic coasts by its high pectoral count (16-17 ; cf. 14-15), smaller jaw teeth and very short and blunt snout. Steindachner 's description and figure (see Plate 3c) are adequate and the provision of a neotype is unnecessary at present. 42 P. J. P. WHITEHEAD TABLE 2 Alphabetical list of nominal species referable to the genus Ilisha. Numbers preceding name refer to senior synonyms listed in key (p. 21). 7. abnormis (Ilisha) Richardson, 1846, Rept. Ichthyol. China Japan : 306 (Whitehead, 1966 : 32). 7. affinis (Clupea) Gray, 1830, Illustr. Ind. Zool., 1 : pi. 96 (2) (Whitehead, ig67a : 119). 3. africana (Clupea} Bloch, 1795, Naturg. Aus. Fische, 9 : 45, pi. 407 (Whitehead, 1969!} : 268) I. amazonicum (Pseudochirocentrodon) Miranda-Ribeiro, 1923, Publ. Comm. Linhas Telegr. Estrat. Amazonas, No. 58 : 8 (Myers, 1950 ; Hildebrand, 1964 : 241). 4. amblyuropterus (Pellona) Bleeker, 1852, Verh. Bat. Gen., 24 : 21 (Whitehead et al., 1966 : 94). i. apapae (Ilisha} Hildebrand, 1948, Smithson. misc. Coll., 110 (9) : 3, fig. 2 (Myers, 1950). 8. brachysoma (Pellona) Bleeker, 1852, Verh. Bat. Gen., 24 : 22 (Whitehead et al., 1966 : 100 ; Whitehead, I967a : 116). 7. chinensis (Pristigaster) Basilewski, 1855, Nouv. Mem. Soc. Nat., Moscow, 10 : 243 (Fowler, 1941 : 662). ?8. ditchoa (Pellona} Valenciennes, 1847, Hist. Nat. Poiss., 20 : 313 (Whitehead, i967a : 116). 3. dolloi (Pristigaster) Boulenger, 1902, Proc. zool. Soc. London : 271, pi. 30 (3) (Whitehead, I967a : 112). 9. dussumieri (Pellona) Valenciennes, 1847, Hist. Nat. Poiss., 20 : 316, pi. 596 (Whitehead, 19673. : 113). 7. elongata (Alosa) Bennett, 1830, Mem. Life of Raffles : 691 (Whitehead, 19673. : 119). 6.filigera (Pellona) Valenciennes, 1847, Hist. Nat. Poiss., 20 : 322 (Whitehead 19673, : 117). 2.furthii (Pellona) Steindachner, 1875, Sitzb. K. Akad. Wiss. Wien, 70 : 388 (Hildebrand, 1946 : 91 ; see also p. 22). 3. gabonica (Pellona) Dumeril, 1858, Arch. Mus. Hist, nat., 10 : 259, pi. 3 (3) (Whitehead, 19673. : 112). 7. grayana (Pellona) Valenciennes, 1847, Hist. Nat. Poiss., 20 : 315 (Whitehead, i967a : 119). 8. indicus (Platygaster) Swainson, 1839, Nat. Hist. Anim., 2 : 294 (Whitehead, 19673. : 114, 117). 3. iserti (Pellona) Valenciennes, 1847, Hist. Nat. Poiss., 20 : 307 (Whitehead, ig67a : 112). i. iquitensis (Ilisha) Nakashima, 1941, Boll. Mus. Hist. Nat. "Javier Prado ", 5 (16) : 66, fig. (Myers, 1950). 9. kampeni (Pellona) Weber & De Beaufort, 1913, Fish. Indo-Austr. Arch., 2: 87. 7. leschenaulti (Pellona) Valenciennes, 1847, Hist. Nat. Poiss., 20 : 311 (Whitehead, 19673. : 118). STEINDACHNER'S CLUPEOID FISHES 43 5. macrogaster (Ilisha) Bleeker, 1866, Ned. Tijdschr. Dierk., 3 : 300 (Whitehead et al., 1966 : 98). 9. macrophthalma (Platygaster} Swainson, 1838, Nat. Hist. Anim., 1 : 278 (nomen oblitum — Whitehead, I967a : 115). 9. megalopterus (Platygaster} Swainson, 1839, Nat. Hist. Anim., 2 : 294 (Whitehead et al., 1966 : 103 ; Whitehead, igGya : 114). 3. melanota (Ilisha} Derscheid, 1924, Rev. Zool. Africaine, Bruxelles, 12 : 278 (Whitehead, 19673. : 112). 8. melastoma (Clupea) Schneider, 1801, Syst. Ich. Block. : 427 (nomen oblitum— Whitehead, 19690 : 270). 8. micropus (Pellona) Valenciennes, 1847, Hist. Nat. Poiss., 20 ; 320 (Whitehead, ig67a : 116). ?8. motius (Clupanodon} Hamilton-Buchanan, 1822, Fishes of Ganges : 251, 383 (Whitehead, 19673 : 115). 7. novacula (Pellona} Valenciennes, 1847, Hist. Nat. Poiss., 20 : 319 (Whitehead, I967a : 121). 2. panamensis (Pellona} Steindachner, i8j5,Sitzb. K. Akad. Wiss. Wien, 70 (i) : 389 (see p. 24). 8. parva (Platygaster} Swainson, 1839, Nat. Hist. Anim., 2 : 294 (on Gray, 1834, Illustr. Ind. Zool., 2 : pi. 109 (3), Clupea motius}. 4. pristigastroides (Pellona} Bleeker, 1852, Verh. Bat. Gen., 24 : 20 (Whitehead et al., 1966 : 93). 9. russellii (Pellona} Bleeker, 1852, Nat. Tijdschr. Ned. Ind., 3 : 72 (Whitehead et al., 1966 : 101). 7. schlegelii (Pellona} Bleeker, 1854, Nat. Tijdschr. Ned. Ind., 6 : 418 (Whitehead et al., 1966 : 99). 7. sinensis (Pristigaster} Sauvage, 1881, Bull. Soc. Philom. Paris, (7) 5 : 107 (White- head, I967a : 119). 4. sladeni (Pellona} Day, 1869, Proc. zool. Soc. London : 623. 8. verticalis (Platygaster} Swainson, 1838, Nat. Hist. Anim., 1 : 278 (Whitehead, 19673. : 116). 7. vimbella (Pellona} Valenciennes, 1847, Hist. Nat. Poiss., 20 : 317 (Whitehead, i967a : 120). 6. xanthopterus (Pellona) Bleeker, 1851, Nat. Tijdschr. Ned. Ind., 2 : 439 (Whitehead et al., 1966 : 96). 44 P. J- P. WHITEHEAD TABLE 3 Gillraker counts in specimens of Pellona castelnaeana and P. flavipinnis. P. castelnaeana (Amazon basin) S.L. gillrakers on lower arch SYNTYPE MNHN.3705 380 14 SYNTYPE MNHN.3706 266 13 BMNH. 1925. 10. 28. 3 470 13 BMNH. 1925. 10. 28.4 365 12 P. altamazonica (fide Hildebrand, 1964) 220-275 12-13 Guyana, Rupununi savannas.) 363 13 BMNH. 1969.7.17.98 P. flavipinnis (Amazon basin) BMNH. 1897. 7. 17. 19 (Amazon mouth) 345 23 NMV.iioi (Teffe) 340 26 BMNH. 1929. 11.18.2 305 25 BMNH. 1869. 5. 21. 51 260 28 NMV.U2 (Iquitos, Peru) 118 28 105 29 (Guyana, Surinam) BMNH. 1934.9. 12. 2 (head only, 125 mm) Approx. 500 mm 24 ZMA. (Marowijne R., Surinam) 438 23 408 23 37° 25 BMNH. 1932.11.10.1 300 25 RMNH. (Surinam) 253 25 RMNH. 3344 (Surinam) 267 25 248 25 BMNH. 1843.6.22.107 240 25 RMNH. 1675 (Surinam) 202 24 BMNH. 1936.5.6.1 197 25 BMNH. 1961.8.31.60 176 24 RMNH. (Surinam) 158 25 (Argentina) BMNH. 1965. 9. 8. 28 (Rosario) 250 31 BMNH. 1878. 5.16. 2 (Buenos Aires) 130 29 BMNH. 1965. 9. 8. 22-27 (Rosario) 114-145 29-31 BMNH. 1969.11.25.146 78 29 STEINDACHNER'S CLUPEOID FISHES 45 REFERENCES BERRY, F. H. 1964. Review and emendation of : Family Clupeidae by Samuel F. Hilde- brand. Copeia, No. 4 : 720-730. BERTIN, L. 1943. Revue critique des Dussumierids actuels et fossiles. Description d'un genre nouveau. Bull. Inst. oceanogr. Monaco, No. 853 : 1-32. BLACKBURN, M. 1949. Age, rate of growth and general life-history of the Australian pilchard (Sardinops neopilchardus) in New South Wales waters. Bull. C.S.I.R., Melbourne, No. 242 : 1-86. CHABANAUD, P. 1926. Sur les clupeides du genre Sardinops Antipa et de divers genres voisins. Bull. Soc. zool. France, 51 : 156-163. CHAN, W. L. 1965. A systematic revision of the Indo-Pacific clupeid fishes of the genus Sardinella (Family Clupeidae). Jap. J. Ichthyol., 12 (3-6) : 104-118 ; Ibid, 13 (1-3) : 1-39- DE PLAZA, M. L. F. 1962. Une nueva especie de Anchoa de las aguas argentinas, Lycen- graulis simulator (Pisces : Engraulidae) . Physis, 23 (64) : 1-9. FOWLER, H. W. 1941- Contributions to the biology of the Philippine Archipelago and adjacent regions. Bull. U.S. natl. Mus., 13 (100) : 1-879. GUNTHER, A. C. L. G. 1868. Catalogue of the fishes in the British Museum, 7, London, 512 pp. — 1909. Fische der Siidsee, 8. /. Mus. Godeffroy, 16 : 261-388. HILDEBRAND, S. F. 1943- A review of the American anchovies (Family Engraulidae). Bull. Bingham oceanogr. Coll., 8 (2) : 1-165. - 1946. A descriptive catalog of the shore fishes of Peru. Bull. U.S. natl. Mus., No. 189 : 1-530- — 1948. A new genus and five new species of American fishes. Smithson. Miser Coll., 110 (9) : 1-15- 1964. Fishes of the Western North Atlantic, part 3. Sears Foundation for Marine Research, New Haven. Memoir i, 630 pp. JORDAN, D. S. 1895. The fishes of Sinaloa. Proc. Calif. Acad. Sci., (2) 5 : 377-514. JORDAN, D. S., EVERMANN, B. W. & CLARK, H. W. 1930. Check list of the fishes and fish-like vertebrates of North and Middle America, north of the boundary of Venezuela and Columbia. Appendix to Kept. U.S. Comm. Fisheries (1928) (2) : 1-670. KAHSBAUER, P. 1959. Intendant Dr. Franz Steindachner, sein Leben und Werk. Annln. naturh. Mus. Wien, 63 : 1-30. LIMA, H. DE H. 1966. Sobre a occorencia de Sardinella anchovia Cuvier & Valenciennes, 1847 no nordeste Brasiliero. Archos Est. Biol. mar. Univ. Ceara, 6 (i) : 67-69. LONGLEY, W. H. & HILDEBRAND, S. F. 1941. Systematic catalogue of the fishes of Tortugas, Florida. Paps. Tortugas Lab., 34 (Carnegie Inst. Publ. No. 533) : 1-331. MANN, F. G. 1954. Vida de los peces en aguas chilenas. Santiago, Chile, 339 pp. MEEK, S. E. & HILDEBRAND, S. F. 1923. The marine fishes of Panama. Field Mus. nat. Hist. (Zool. Ser.), 15 (i) : 1-330. MYERS, G. S. 1950. Systematic notes on some Amazonian clupeid fishes of the genus Ilisha. Copeia, No. i : 63-64. NORMAN, J. R. i923a. A revision of the clupeid fishes of the genus Ilisha and related genera. Ann. Mag. nat. Hist., (9) 11 : 1-22. i923b. A new species of Neosteus from Rio de Janeiro, with notes on the American species of Ilisha and Neosteus. Ann. Mag. nat. Hist., (9) 12 : 593-4. REGAN, C. T. 1916. British fishes of the subfamily Clupeinae and related species in other seas. Ann. Mag. nat. Hist., (8) 18 : 1-19. - 1917. A revision of the clupeid fishes of the genera Sardinella, Harengula, &c. Ibid : 377-395- RIBEIRO, A. DE M. 1923. Historia natural, Zoologia : Peixes (excl. Characinidae) . Comm. Linhas Telegr. Estr. Matto-Grosso Amazonas, Publ. No. 58, Annexo No. 5 : 1-15. RIVAS, L. R. 1964. Genus Harengula Cuvier and Valenciennes, 1847. In Fishes of the 46 P. J. P. WHITEHEAD Western North Atlantic, part 3. Sears Foundation for Marine Research, New Haven, Memoir i, 630 pp. ROSSIGNOL, M. 1955. Premieres observations sur la biologic des sardinelles dans la region de Pointe Noire (Sardinella eba Val., S. aurita Val.). Rapp. Cons. Explor. Mer., 137 : 17-21. SCHMELTZ, J. D. E. 1869. Museum Godeffroy, Catalog IV, 139 pp. SCHULTZ, L. P. & WELLANDER, A. D. 1953. Fishes of the Marshall and Marianas Islands. Butt. U.S. nail. Mus., No. 202 : 1-685. STEINDACHNER, F. 1882. Beitrage zur Kenntniss der Fische afrika's (II). Denkschr. Akad. Wiss. Wien., 45 (i) : 1-18. SVETOVIDOV, A. N. 1952. Fauna of the U.S.S.R. — Fishes, 2 (i) (N.S. No. 48). [English ver- sion 1963, Jerusalem, 428 pp.]. - 1964. Systematics of the North American anadromous clupeoid fishes of the genera Alosa, Caspialosa and Pomolobus. Copeia, No. i : 118-130. VALENCIENNES, A. 1847. Histoire naturelle des poissons, 20, Paris, 472 pp. - 1848. Ibid. 21, Paris, 536 pp. WHITEHEAD, P. J. P. 1962. A revision of the recent round herrings (Pisces : Dussumieriidae). Bull. Br. Mus. nat. Hist. (Zool.), 10 (6) : 305-380. — ig64a. A redescription of the holotype of Clupalosa bulan Bleeker, and notes on the genera Herklotsichthys, Sardinella and Escualosa (Pisces : Clupeidae). Ann. Mag. nat. Hist., (13) 7 : 33-47- - I964b. New data extending the range of the bipolar antitropical anchovy genus Engraulis into the tropics. Zool. Zh., 43 (6) : 879-888 (in Russian). I9&5a. A review of the elopoid and clupeoid fishes of the Red Sea region. Bull. Br. Mus. nat. Hist. (Zool.), 12 (7) : 225-281. — i965b. A preliminary revision of the Indo-Pacific Alosinae (Pisces : Clupeidae). Bull. Br. Mus. nat. Hist. (Zool.), 12 (4) : 115-156. 1966. The elopoid and clupeoid fishes in Richardson's 'Ichthyology of the seas of China and Japan, 1846.' Butt. Br. Mus. nat. Hist. (Zool.), 14 (2) : 15-44. - ig6ja. The clupeoid fishes described by Lacepede, Cuvier and Valenciennes. Bull. Br. Mus. nat. Hist. (Zool.), Suppl. 2 : 1-180. - 1967^ The West African shad, Ethmalosa fimbriata (Bowdich, 1825) : synonymy, neo- type. /. nat. Hist., 4 : 585 — 593. 1968. A new genus for the South American clupeid fish Lile platana Regan. /. nat. Hist., 2 : 477-486. — i96ga. The clupeoid fishes of Malaya. /. mar. biol. Assn. India, 9 (2) : 223-280. — ig69b. The clupeoid fishes described by Bloch and Schneider. Bull. Br. Mus. nat. Hist. (Zool.), 17 (7) : 261-279. WHITEHEAD, P. J. P., BOESMAN, M. & WHEELER, A. C. 1966. The types of Bleeker's Indo- Pacific elopoid and clupeoid fishes. Zool. Verhandl. Leiden, No 84 : 1-152. WHITLEY, G. P. 1940. Illustrations of some Australian fishes. Austr. Zool., 9 (4) : 397-428. P. J. P. WHITEHEAD, B.A. Department of Zoology BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON, S.W.7 D PLATE i a. Alausa alburnus Kner & Steind. ( = Spratelloides delicatulus] b. Pellonula bahiensis Steind. ( — Rhinosardinia bahiensis) c. Alausa fimbriata Kner & Steind. ( = Sardinops sagax sagax) Bull. Br. Mus. nat. Hist. (Zool.) 20, i PLATE i Q PLATE 2 a. Clupea setosa Steind. ( = Ethmalosa fimbriata) b. Clupea notacanthoides Steind. ( = Ethmidium maculatum notacanthoides) Bull. Br. Mus. nat. Hist. (Zool.) 20, i PLATE 2 a PLATE 3 a. Engraulis nasus Kner & Steind. ( — Anchoa nasus) b. Engraulis macrolepidotus Kner & Steind. ( = Anchovia macrolepidota) c. Engraulis poeyi Kner & Steind. ( = Lycengraulis poeyi) Bull. BY. Mus. nat. Hist. (Zool.) 20, i PLATE 3 a Printed in Great Britain by AJden & Mowbray Ltd at the Alden Press, Oxford THE TYPE SPECIMENS OF SIPUNCULA AND ECHIURA DESCRIBED BY J. E. GRAY AND W. BAIRD J IN THE COLLECTIONS OF THE BRITISH MUSEUM (NATURAL HISTORY) MARY E. RICE & A. C. STEPHEN BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 20 No. 2 LONDON: 1970 THE TYPE SPECIMENS OF SIPUNCULA AND ECHIURA DESCRIBED BY J. E. GRAY AND W. BAIRD IN THE COLLECTIONS OF THE BRITISH MUSEUM (NATURAL HISTORY) BY MARY ESTHER RICE National Museum of Natural History, Smithsonian Institution, Washington Vy and ALEXANDER CHARLES STEPHEN Royal Scottish Museum, Edinburgh Pp. 47-72 ; 3 Plates BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 20 No. 2 LONDON : 1970 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1940, is issued in five series corresponding to the Departments of the Museum, and an Historical series. Parts will appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department. This paper is Vol. 20 No. 2 of the zoological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation Bull. Br. Mus. nat. Hist. (Zool.). Trustees of the British Museum (Natural History), 1970 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 9 November, 1970 Price £1.20 THE TYPE SPECIMENS OF SIPUNCULA AND ECHIURA DESCRIBED BY J. E. GRAY AND W. BAIRD IN THE COLLECTIONS OF THE BRITISH MUSEUM (NATURAL HISTORY) By MARY E. RICE & A. C. STEPHEN SYNOPSIS Type specimens of Sipuncula and Echiura of Gray (1828) and Baird (1868) in the collections of the British Museum (Natural History) were re-examined by the authors. Identifications were checked and, in most instances, the specimens were redescribed. Of the 23 type specimens, the specific names of the following were shown to be senior subjective synonyms of currently accepted names : Siphunculus arcuatus Gray 1828 [ = Phascolosoma lurco (Selenka and de Man 1883)], Phascolosoma perlucens Baird 1868 [ = Phascolosoma dentigerum (Selenka and de Man 1883)], Themiste lageniformis Baird 1868 [ = Themiste signifer (Selenka and de Man 1883)], and Aspidosiphon jukesii Baird 1873 [ = Aspidosiphon corallicolus Sluiter 1902]. Five were shown to have currently accepted valid names, one was a nomen dubium, and the remainder were junior synonyms to currently used names. INTRODUCTION DURING the course of the preparation of a monograph on the Sipuncula and Echiura, it became necessary to clarify the status of the several species described by Gray (1828) and Baird (1868 and 1873) since many were originally described mainly on external characters. Where these authors' specimens are still preserved in the col- lections of the British Museum (Natural History), it has been possible to check their identifications and, in most instances, to redescribe them. Because of the historic value of the specimens, only a minimum of dissection and manipulation was attempted ; hence complete redescriptions are not always provided in this report. This study was initiated by the late A. C. Stephen of the Royal Scottish Museum and, after Dr. Stephen's death in 1966, it was completed by Mary E. Rice of the Smithsonian Institution. Thanks are due to the Trustees, to Dr. J. P. Harding, Keeper of Zoology, and to Mr. R. W. Sims, Head of the Annelida Section, British Museum (Natural History) for permission to examine the specimens. Mrs. Carolyn Gast, scientific illustrator at the Smithsonian Institution, is gratefully acknowledged for her illustrations of the type specimens. THE GRAY TYPES Gray (1828 : 8, pi. 6, figs, i, 4, 4a) described a series of sipunculans from the collec- tions of the British Museum (Natural History) under the heading " Radiata ", 50 MARY E. RICE & A. C. STEPHEN family Siphunculidae. He included six species, four of which were described as new. These were : Siphunculus dentallii, Siphunculus tuberculatus , Siphunculus arcuatus, and Themiste hennahi. Gray's brief diagnoses were in Latin, followed by a few comments in English, and they pertained only to non-specific external characters. Consequently, even though Gray's type specimens remained extant in the British Museum (Natural History), they were ignored by most subsequent authors. A re-examination of the specimens shows that the species names of 5. arcuatus and T. hennahi are valid, whereas 5. tuberculatus, demonstrated by Baird (1868) to be a junior primary homonym and re- named by him as Phascolosoma grayi, is here synonymized with Phascolosoma noduli- ferum Stimpson 1855. Specimens of 5. dentalii are missing from the collection and apparently lost. Siphunculus arcuatus Gray, 1828 Siphunculus arcuatus Gray, 1828, p. 8. Phascolosoma arcuatum : Baird, 1868, p. 88. HOLOTYPE : Reg. No. 1965. 25. 2 TYPE LOCALITY : India. Coll. Hardwicke. DESCRIPTION : The specimen was preserved in a curved position with the introvert entirely retracted. It is in good condition with most of the internal organs well preserved, possibly because an incision had been made at one time in the body wall. The trunk measures approximately 100 mm in length and 20 mm at its maximum width. The retracted introvert is slightly longer than the trunk and is coiled and twisted within the body cavity. The basic colouration of the trunk is pale brown, although in the anterior third of the trunk there is an area of reddish-brown pigmenta- tion. The papillae, distributed over the entire trunk, stand out as dark brown spots against the lighter background ; they are largest and most concentrated at the anter- ior and posterior extremities of the trunk (Figure 3, 3a-3c). The anterior and pos- terior papillae are pyrimidal in shape, the largest measuring 0-8 mm in width and 0-5 mm in height in the posterior region ; the middle papillae are lower and more rounded, attaining a size of 0-5 mm in width and 0-24 mm in height. The platelets of the posterior and anterior papillae are dense and compact and in some cases more darkly pigmented toward the center of the papilla, whereas those of the middle papillae are more dispersed and evenly coloured (Figure 2). Surrounding the central opening of the papilla the platelets are smaller ; otherwise there is no obvious gradient in size of platelets. The hooks are unidentate, strongly curved, with a clear central streak which is markedly wider at its basal extremity (Figure 2). The basal portion of the hook is diaphanous in nature and very much thinner than the remainder of the structure. Determination of the number of rows of hooks and number of tentacles would have required excessive manipulation of the retracted introvert of this historically valuable specimen ; hence, no observations were made on these characters. In distended areas of the body wall in the posterior half of the trunk, the cuticle is inflated into a series of small, thin-walled sacs (Figure 4). Upon dissection of the animal these sacs SIPUNCULA AND ECHIURA 51 appear to be vesicular expansions of the coelomic cavity into the body wall, extending through spaces between longitudinal and circular muscles. The integument covering the vesicles is markedly thinner than that of the remainder of the trunk. Circular muscles, as well as longitudinal muscles, are divided into bundles. The longitudinal bundles show little anastomosis, numbering 18 at the level of the nephri- diopores and 19 in the region of the posterior nephridia. The bundles of the circular muscles are smaller, more numerous, and less distinctive with frequent anastomoses. Two posterior retractor muscles originate in the posterior sixth of the trunk from longitudinal muscle bundles 2 and 3 and the two muscles soon join to form a single posterior retractor (Figure i). The anterior retractors originate on either side of the ventral nerve cord, from longitudinal muscle bundle number i at the anterior end of the posterior quarter of the trunk near the level of the union of the posterior retrac- tors. The anterior retractors soon join the posterior retractor to form a single retractor muscle (Figure i). The spindle muscle is attached at the posterior extremity of the trunk and after coursing anteriorly through the center of the intestinal coil and along the rectum, it attaches to the body wall immediately anteriorly to the anus. Prominent wing muscles fasten the anterior rectum to the body wall. Intestinal coils are numerous, but the exact number is difficult to ascertain because the coiling is somewhat erratic and parts of the gut are not well preserved. The nephridia are approximately one-half as long as the trunk and are attached to the body wall for nearly their entire length. The nephridiopores open slightly anter- ior to the anus. REMARKS : Gray's holotype of Siphunculus arcuatus corresponds with descriptions of Phascolosoma lurco (Selenka and de Man 1883, p. 61-63) in the following significant taxonomic characters : structure of hooks on introvert, form and distribution of papillae, fusion of four retractors into a single, long retractor muscle, and the number of longitudinal muscles. In Phascolosoma lurco, as in the holotype of Siphunculus arcuatus, the origin of the anterior retractor muscles is ventral to that of the posterior retractors rather than the more common arrangement for sipunculans in which the posterior muscles originate ventral to the anteriors. One exception for Phascolosoma lurco was found in speci- mens examined by Lanchester (1905) in which both pairs of retractors originated from the same longitudinal line. The point of fusion of the four retractors in Phascolosoma lurco has been described differently by various authors. Selenka's figure depicts a fusion of the left anterior and posterior retractor muscles and a separate fusion of the two right retractors, resulting in one left and one right retractor which, after a short distance, unite to form a single muscle. Both Lanchester (1905) and Edmonds (1956), on the other hand, report that the four muscles fuse at about the same level to form one long retractor muscle. Siphunculus arcuatus presents a further varia- tion : the two posterior muscles fuse to form one central muscle which courses anter- iorly for a short distance and is then joined on either side by the left and right anterior muscles. In Phascolosoma lurco as in Siphunculus arcuatus, the circular musculature shows a propensity for separation into bundles, although the bundles are not as widely spaced 52 MARY E. RICE & A. C. STEPHEN nor as prominent or regular as those of the longitudinal muscles (Edmonds 1956, Selenka, de Man and Billow 1883). In his description of Phascolosoma lurco, Selenka mentions small inflated areas in the body wall which may be comparable to the coelomic vesicles found in Gray's holotype. In a synonymy of the two names, the specific name arcuatus has priority over lurco. Siphunculus dentalii Gray, 1828 Siphunculus dentalii Gray 1828, p. 8 ; Johnston, 1833, p. 233-235, fig. 25. TYPE LOCALITY : Coast of Yorkshire, in Dentalium. Coll. Clift. This specimen is missing from the collection. It was not illustrated in Gray's report and his description which lists only general external characters is inadequate for a determination of the species involved. The locality of the specimen and its habitat in the shell of a Dentalium, suggest that it may have been Phascolion strombus (Montagu). However, Gray also described and figured a specimen of Phascolion strombus which he designated as Siphunculus strombus Mont. ; thus, it is obvious that he did not consider the two specimens to be the same, although the differences in his descriptions are non-specific, related mainly to shape and size of trunk and introvert. In 1833, Johnston figured 5. dentalii and elaborated on Gray's description, but he does not indicate whether he examined Gray's type specimen. Later authors (Selenka, de Man, and Biilow 1883, Gerould 1913, tenBroeke 1929) have placed 5. dentalii Gray in synonymy with Phascolion strombus (Montagu). Siphunculus tuberculatus Gray, 1828 Siphunculus tuberculatus Gray, 1828, p. 8 (non Siphunculus tuberculatus de Blainville, 1827). Phascolosoma grayi (nom. nov.) Baird, 1868, p. 88. HOLOTYPE : Reg. No. 1965.25.4 TYPE LOCALITY : Unknown Gray (1828) described this specimen, to which he gave the name Siphunculus tuberculatus, as follows : " The body is conical and attenuated behind ; the trunk tubercular at the base, and nearly smooth at the apex. Length of the body i| inch, its breadth £ inch ; length of the trunk \ inch ". In 1868 Baird pointed out that the specific name tuberculatus had been preoccupied by de Blainville in 1827. On re- examining Gray's specimen he concluded that it was different from de Blainville's species and renamed it Phascolosoma grayi, after Gray, who first defined the species. The type is now in rather poor condition. It had been previously dissected and most of the internal organs have been lost. DESCRIPTION : The length of the body with fully extended introvert is approxi- mately 60 mm, the maximum width 5 mm. The length of the introvert is slightly less than that of the trunk. The overall colour is a pale greyish yellow, somewhat darker at the base of the introvert and posterior end. The base of the introvert, the preanal region and the posterior quarter of the trunk are marked by a dense SIPUNCULA AND ECHIURA 53 accumulation of prominent, rounded, dome shaped papillae (Figure u), the largest of which measures approximately 0-32 mm in diameter and 0-28 mm in height. In the middle region of the trunk the papillae are smaller, measuring as much as 0-20 mm in diameter and 0-08 mm high, and although distributed in a regular pattern they are more widely separated than at the extremities of the trunk. Platelets are arranged in a distinctive pattern in all papillae : a ring of dark brown platelets surrounds a clear central area and light-colored widely spaced platelets cover the remainder of the papilla (Figure 10). Roughly 20 rows of dark brown hooks encircle the anterior quarter of the introvert followed by numerous irregularly placed hooks. Simple in structure, the hooks are bent terminally and show a single clear central streak (Figure 9). Tentacles, although retracted, had been exposed by a previous dissection and 12 long filiform processes were counted. The skin of the middle trunk is characterized by transverse wrinkling. Longi- tudinal wrinkling, weak and irregular in the middle trunk, is more pronounced in preanal and posterior regions. Internally only the rectum, nerve cord, and nephridia remain in this incompletely preserved specimen. The nephridiopores and anus open at the same level. A broad wing muscle attaches the rectum to the body wall and the spindle muscle runs along the length of the rectum, inserting on the body wall immediately anterior to the anus. Only one nephridium, the left, is intact and it is attached for most of its length to the body wall with only the posterior end free. The longitudinal musculature is arranged in bundles which undergo considerable anastomosis, but at the level of the nephridio- pores 25 bundles can be counted. The retractor muscles have been broken off and only remnants remain attached in the anterior introvert. REMARKS : The characters which are still recognizable in this incomplete specimen agree closely with those of Phascolosoma noduliferum Stimpson 1855, as enumerated in a recent review of the species by Edmonds (1956). The number of muscle bundles falls within the range for P. noduliferum and the structure and distribution of the distinctive hooks and papillae are identical. Although Baird listed Phascolosoma noduliferum in his monograph (1868), he failed to recognize the similarities between this species and the specimen designated by Gray as Siphunculus tuber culatus. Hence, after indicating that Gray's name was pre-occupied, Baird provided a new name, Phascolosoma grayi, whereas it appears that he should have synonymized the junior homonym with P. noduliferum Stimpson. Themiste hennahi Gray, 1828 Themiste hennahi Gray, 1828, p. 8 ; Baird, 1868, p. 98 ; Stephen, 1964, p. 458 ; Amor, in press. LECTOTYPE : Reg. No. 1965.16.1. PARALECTOTYPES : Reg. No. 1965.16.2/5 TYPE LOCALITY : Peru. Coll. Rev. Hennah. Stephen (1964), in a reclassification of the Sipuncula, demonstrated that the name Themiste Gray, 1828, was the senior synonym of the genus Dendrostomum Grube, 1858. Gray figured the tentacular crown of Themiste hennahi, the type species of Themiste 54 MARY E. RICE & A. C. STEPHEN by monotypy, and clearly showed the dendritic nature of the tentacles but this paper was overlooked by Grube who erected the genus Dendrostomum for species with dendritic tentacles. Gray's original description of Themiste hennahi, quoted in full by Stephen as it had been printed in a thinly distributed publication of which few copies survive, was brief and inadequate by modern standards ; hence, a complete redescription of the specimens to which Gray gave the name Themiste hennahi is presented here. From the five specimens still intact at the British Museum (Natural History), Stephen (1964) designated one lectotype and four paralectotypes. DESCRIPTION : The lectotype designated by Stephen (1964) is a fully extended specimen, measuring 70 mm in length to the base of the tentacles with a maximum width of approximately 15 mm (Figure 8). The introvert is one-fifth the length of the body and it is characterized by a relatively short smooth zone (5 mm) adjacent to the tentacular crown. The cuticle of the posterior introvert has become detached from the underlying epidermis, exposing the protruding canals of the epidermal glands (Figure 8a), and forming a wrinkled mass at the base of the introvert. No hooks or spines are apparent on the introvert. The predominant colour of the body is a pale grey, whereas that of the tentacles and smooth zone is light tan. The well-extended dendritic tentacles arise from six primary stems (Figure 7). Between the bases of the tentacular stems are the six lappets or lips, membranous crescentic folds which form the margin of the mouth and demarcate the proximal boundaries of the six primary food grooves leading into the mouth from the tentacular stems. The dorsal lappet is distinguished by its large size and its proximity to the nuchal organ. The tentacular crown is asymmetric, the lateral tentacles exceeding both the dorsals and ventrals in length. The primary food grooves of the lateral stems are also longer, bifurcating at a greater distance from the mouth than the ventral and dorsal grooves. The openings of the epidermal glands are distributed in a regular pattern over the surface of the trunk ; in the anal region the openings are in the form of slits and are situated on oblong elevations of the skin (Figure 8b) . More posteriorly the openings approach a spherical shape ; in the middle of the trunk they are situated between transverse furrows of the skin, but no distinctive elevations are apparent (Figure 8c). In the posterior portion of the trunk the skin is marked by longitudinal as well as transverse furrows, resulting in small irregular rectangles within each of which is enclosed a gland opening. At the posterior extremity of the body the openings are elevated on dome-shaped swellings of the skin (Figure 8d). The position of the nephridiopores is slightly posteriorly to that of the anus. Since the lectotype had not been dissected, it was left intact and the internal anatomy of this specimen was not studied. However, of the four paralectotypes, two had been previously dissected and the better preserved and more clearly dissected specimen was selected for the description of internal anatomy which follows (Figure 6) . The introvert of this paralectotype was retracted and the length of the body with- out introvert was approximately 37 mm. The musculature of the body wall is smooth and iridescent. A pair of wide, thickened retractor muscles originate on either side of the ventral nerve cord at the beginning of the posterior third of the trunk. The spindle muscle is not attached SIPUNCULA AND ECHIURA 55 posteriorly, but emerges anteriorly from the intestinal coil and, after giving off a short branch to the caecum, continues along the rectum, attaching to the body wall slightly posteriorly to the level of the anus. Three fixing muscles were observed (Figure 6). Fi arises on the left side of the body in the anterior third of the trunk in a ventrolateral position and inserts on the oesophagus in the region where the contractile vessel breaks up into many blind vessels. The F2 unfortunately had been broken in this specimen and its point of insertion was not determined ; its origin is in a ventrolateral position on the left side of the body slightly posteriorly to the origin of Fj. The fixing muscle F3 is a short thick muscle which attaches the last ascending intestinal coil to the dorsal body wall. The oesophagus is long, narrow, and thin-walled and, in this specimen it is looped under the right retractor (Figure 6). Running along the dorsal side of the oeso- phagus is the contractile vessel, enormously distended anteriorly. It is attached to the oesophagus for approximately one-fifth the length of the latter and at its posterior end it breaks up into many blind vessels or villi which ramify throughout the body cavity. Resembling strings of beads, these processes of the contractile vessel are marked by globular enlargements connected by thin, narrow, sometimes coiled strands. The intestine is wound into numerous coils (approximately 40 single coils), difficult to count with accuracy because of irregular winding and poor preserva- tion in some parts. The wall of the gut is very thin and the ascending gut is filled with fine sand or mud particles. The beginning of the rectum is marked by the presence of a small caecum and in the anal region the rectum is attached to the body wall by broad wing muscles. Two nephridia, more than half the length of the trunk, hang freely in the body cavity. The anterior portions of the nephridia are swollen and distended ; the nephrostomes are small and simple. Nephridiopores open slightly posteriorly to the anus. In addition to the lectotype and paralectotype described above, there are three remaining paralectotypes of Themiste hennahi. One of these is a small specimen which had been dissected ; its tentacles are partially extended and the trunk, without the introvert, measures approximately 20 mm in length. In the other two paralecto- types the introvert is retracted and the approximate measurements of the lengths of the trunks are 45 mm and 40 mm. REMARKS : The number of tentacle stems was recorded originally as five (Gray 1828). Stephen (1964), looking at the same specimen which he designated as the lectotype, reported Gray's count of tentacles to be in error, stating that "... there are only four tentacle stems, one shows a false dichotomy which misled Gray into thinking that there were five ". A re-examination of this specimen by one of the present authors (Rice) has shown, in contradiction to the two previous reports, that there are six tentacular stems with six distinctive primary food grooves. The oral view of the tentacular crown illustrated in Figure 7 clearly shows the six primary stems, food grooves and lips. Gray figured a lateral view of the anterior end of this same specimen, but his view does not clearly delineate the number of tentacles. Perhaps the past discrepancy has been due in part to a failure to regard the number and position of lips. Moreover, the variation in length, size, and branching of the 56 MARY E. RICE & A. C. STEPHEN tentacles and the differing lengths of the primary food grooves of the specimen prove to be confusing in any attempt to count tentacles. Another possible source of error in the interpretation of tentacular arrangement is the absence of one of the tentacles from the right ventral tentacular stem ; apparently it had been broken off at some time just beyond the point of the first bifurcation of the primary stem, and only the base of the tentacle and the beginning of the secondary food groove remain (Figure 7)- The two species Dendrostomum peruvianum Collin, 1892, and Themiste hennahi Gray were synonymized by Stephen in 1964. D. peruvianum has been recorded in the literature by two authors in addition to Collin (1892) : Fischer (1914) and Wesenberg-Lund (1955). As in the case of T. hennahi, reports of the number of tentacles of D. peruvianum are found to vary. Fischer (1914) reported five tentacle stems, whereas in the original description of the species Collin mentioned only four. Collin (1892) described slightly raised papillae in the middle and posterior body of Dendrostomum peruvianum ; Fischer denied the presence of typical papillae, but nevertheless described dome-shaped elevations in the posterior body onto which the canals of the skin glands opened. Wesenberg-Lund (1955) reported an absence of projecting papillae, but described instead low circular papillae lying between the wrinkles of skin. In the Gray lectotype projecting papillae were not found in the middle of the trunk but in the posterior and anal regions the glands opened on elevated protuberances of the skin. (Esophageal protuberances described by Collin were not seen by Wesenberg-Lund on the specimen which she identified as Dendrostomum peruvianum, nor were they observed in this study on the Gray paralectotype. As in Fischer's specimen of D. peruvianum, the caecum of the paralectotype of T. hennahi is found at the beginning of the rectum, not on the penultimate intestinal coil as described by Collin. Also varying from Collin's description, the F3 fixing muscle of the paralectotype is attached to the last intestinal coil rather than the penultimate, and an F4 fixing muscle is absent. In other characters, published descriptions of Dendrostomum peruvianum agree essentially with the Gray specimens. The beaded structure of the villi of the con- tractile vessel, clearly evident in the Gray paralectotype, is described as characteristic of D. peruvianum. Themiste hennahi also shows many similarities to the species described as Dendro- stomum zostericolum Chamberlin 1919 and Dendrostomum schmitti Fisher 1952. The possibility that these may all represent a single species remains to be determined by future studies. THE BAIRD TYPES In 1868 Baird published his monograph on the species of worms belonging to the subclass Gephyrea in the collections of the British Museum. In this he listed 142 species, including 18 descriptions of new species, 17 of which were sipunculans and one a priapulid. Later in a short paper published in 1873, he erected two additional species, Aspidosiphon jukesii and Echi^^rusfarcimen. Baird's descriptions were short and based solely on external characters ; thus, they were inadequate for recognition SIPUNCULA AND ECHIURA 57 by later workers. In Appendix E of their monograph, Selenka, de Man and Billow listed all of the new species described in 1868 by Baird with the exception of Pseudo- aspidosiphon gracile which they considered in the text. They examined six of Baird's type specimens, redescribing and retaining the names of four, and placing two other names in synonymy. The remainder listed in the appendix they considered to be insufficiently described and for some reason which remains unexplained they did not examine these specimens. No further attention seems to have been given to the specimens until Edmonds (1955, 1961), reporting on the sipunculans of Australia, redescribed three of Baird's types. For this report, all of Baird's type specimens have been re-examined, the validity of the names is reviewed, and, where appropriate, they are relegated to the proper synonymy. Sipunculus aeneus Baird, 1868 Sipunculus aeneus Baird, 1868, p. 81. Siphonosoma australe : Edmonds, 1961, pp. 217-220, 2 figs. HOLOTYPE : Reg. No. 1952 : 10.8. TYPE LOCALITY : New Zealand. Coll. Cuming. This specimen was examined by Edmonds (1961) who redescribed it completely and referred it to Siphonosoma australe (Keferstein 1865). Following Baird's original description and previous to Edmonds' report, the name had appeared in the literature only twice : Selenka, de Man, and Billow (1883) listed it in Appendix E of their monograph and Benham (1903) referred to it as a " species inquirenda ". Sipunculus angasii Baird, 1868 Sipunculus angasii Baird, 1868, p. 80, pi. IX, fig. i. Sipunculus angasi : Edmonds, 1955, pp. 83-86, figs. 1-4. SYNTYPES : Reg. No. 1864 : I2.i3.3a/b. TYPE LOCALITY : Port Lincoln, Spencer Gulf, South Australia. Coll. Angas. In his description, Baird (1868) mentioned only non-specific external characters and the name was not used again in the literature until 1955 when Edmonds identified a large number of specimens from Australia as Sipunculus angasi. At that time Ed- monds re-examined Baird's syntypes which he considered to be juveniles and de- scribed the internal anatomy of one of the specimens. Sipunculus angasi was re- garded by Edmonds as closely allied to S. robustus Keferstein and S. nudus Linnaeus. Sipunculus deformis Baird, 1868 Sipunculus deformis Baird, 1868, p. 80-8 1, pi. ix, fig. 2. Siphonosoma cumanense : Edmonds, 1955, p. 90-92. HOLOTYPE : Reg. No. 1965.25.7. TYPE LOCALITY : Sir Charles Hardy's Island, North Australia. Coll. Brookes. Edmonds in 1955 gave a brief description of the internal anatomy of this specimen and considered it to be Siphonosoma cumanense. The only other time the name 58 MARY E. RICE & A. C. STEPHEN Sipunculus deformis has appeared in the literature subsequent to Baird's description was in Appendix E of the monograph by Selenka, de Man and Billow (1883). Sipunculus eximinoclathratus Baird, 1868 Sipunculus eximio-clathratus Baird, 1868, pp. 81-82. HOLOTYPE : Reg. No. 1965.25.8. TYPE LOCALITY : Philippine Islands. Coll. Cuming. Baird's description was limited to external characters and he did not dissect the specimen. It is still in good condition, although a small part of the body wall had been damaged and the viscera at this point destroyed. A dissection was made by one of the authors (ACS) and a description of the internal anatomy is included below. The specimen was preserved in a curved position with most of the introvert re- tracted. Measuring approximately 70 mm in length, the trunk is contracted in the mid-region but distended into bulbous expansions anteriorly and posteriorly. As pointed out by Baird, the skin is divided into rectangular areas by longitudinal and transverse furrows and the introvert is covered with triangular protuberances which point in a posterior direction. Thirty-three longitudinal muscle bundles were counted in the anterior fifth of the trunk and about 30 in the middle. Four short retractor muscles originate quarter of the length of the trunk from the anterior end. The spindle muscle attaches to the body wall anterior to the anus ; a short distance posterior to the anus the characteris- tic " Buschel " organs occur. A caecum is present at the beginning of the rectum. The frons, or cerebral organ, is a simple flap of tissue with no obvious elaborations. These characters correspond with those of Sipunculus nudus Linnaeus, a widely distributed tropical form. Phascolosoma aethiops Baird, 1868 Phascolosoma aethiops Baird, 1868, p. 90. HOLOTYPE : Reg. No. 1839.12.26.46. TYPE LOCALITY : St. Vincent, West Indies. Coll. Guilding (?). In the monograph of Selenka, de Man, and Biilow (1883) the name Phascolosoma aethiops is listed in Appendix E and followed by the question " 1st ein Dendro- stoma? ". Since there is no reason to believe that Selenka ever examined this specimen, it is probable that his question arises from Baird's description of the tentacles as " short and numerous " and his mistaken reference to " small black spines " on the introvert (Baird 1868). DESCRIPTION : The holotype is in rather fragile condition and seems to have been either wholly or partially desiccated at some time. It was dissected by one of the present authors (ACS) and even with a minimum of interference the tentacular crown became detached. The stout, thick trunk is 25 mm long and 8 mm at its maximum width ; the intro- vert is 5 mm in length without the tentacles. The tentacles are filiform, numerous, SIPUNCULA AND ECHIURA 59 and in a position dorsal to the mouth. Even though the tentacles were described by Baird as short, they measure as much as 3 mm in length which is a considerably greater relative length than that found in the majority of species of Phascolosoma. The light brown skin is covered with numerous contrasting dark brown papillae. On the introvert the papillae are conical, resembling spines, although true spines, as cited by Baird, are lacking. Papillae on the trunk are subcircular and are largest and most numerous on the anterior and posterior extremities. Contrary to the situation in most species of this genus, the largest papillae are located ventrally rather than dorsally. Each papilla is characterized by distinctive dark brown platelets which are evenly dispersed around a clear central area. Individual plate- lets are also scattered over the cuticle among the papillae. The longitudinal muscle bundles show considerable anastomosis ; immediately anteriad to the origin of the retractors they number approximately 20. Four retractor muscles originate at nearly the same level in the middle third of the trunk, although the dorsals are slightly anterior to and somewhat thinner than the ventrals. The dorsal and ventral retractors soon join to form left and right retractors which remain separated for most of their length. The gut is attached posteriorly by a spindle muscle and is comprised of approxi- mately 25 single coils. A single fixing muscle extends from an attachment to the body wall left of the ventral nerve cord in the mid-region of the body to the intestinal coil. Strong wing muscles attach the rectum to the body wall in the region of the anus. A prominent contractile vessel with numerous villi runs along the oesophagus and continues into the beginning of the intestinal coil. Part of the left nephridium is missing, but the right nephridium is three-quarters of the length of the trunk and is attached to the body wall for two-thirds of its length. REMARKS : This specimen is identical to Phascolosoma antillarum Grube and Oersted 1859 as evidenced by similarities in the form and distribution of papillae and platelets, tentacular form and pattern, structure of contractile vessel and villi, relative length and attachment of nephridia, the number of longitudinal muscle bundles and the attachment and union of retractor muscles. Phascolosoma albolineatum Baird, 1868 Phascolosoma albolineatum Baird, 1868, p. 91-92. Phymosoma albolineatum : Selenka and de Man, 1883, pp. 71-72, pi. ix, fig. 128-129. HOLOTYPE : Reg. No. 1925.25.1. TYPE LOCALITY : Philippine Islands. Coll. Cuming. The holotype is still in fairly good condition and had been dissected previously. It is presumed to be the specimen described and figured by Selenka, de Man and Billow (1883), since they state that their description is based on Baird's original specimen. They described it fully and the species remains valid. 6o MARY E. RICE & A. C. STEPHEN Phascolosoma capsiforme Baird, 1868 Phascolosoma capsiforme Baird, 1868, p. 83-84, pi. ix, fig. 3 ; Selenka, de Man, and Billow, 1883, p. 27-28, pi. iv, figs. 38-39. SYNTYPES : Reg. No. 1842.2.24.60/63. TYPE LOCALITY : Falkland Islands. Coll. W. Wright. One of Baird' s specimens was examined by Selenka, de Man, and Billow who gave it a full description, including an account of the internal anatomy and figures of the papillae. Since none of the four syntypes in the Museum had been dissected, it must be presumed that the specimen described by Selenka was not returned or has been lost. The four remaining specimens are in excellent condition. One is partially expanded ; the others are contracted. The characters, as reported by Selenka (1883), agree with those of Golfingia margaritacea Sars, a common species of northern seas and now recorded from a num- ber of localities in the Antarctic. Selenka, de Man, and Biilow noted the close re- semblance between the two species, and several authors have since called it a sub- species of the northern species (Fischer 1896, 1913, Benham 1922, Edmonds 1965). In a description of species from the Ross Sea, Edmonds (1965) reviewed previous reports of the northern and southern forms and concluded that the southern species is properly designated as Golfingia margaritacea capsiformis (Baird) . Phascolosoma fasciatum Baird, 1868 Phascolosoma fasciatum Baird, 1868, p. 89. SYNTYPES : Reg. No. 1849.8.4.18/19. TYPE LOCALITY : Madiera (Azores). Coll. N. Lister. The two syntypes are in good condition and neither has been dissected previously. Baird (1868) characterized the species by the brown bands on the introvert, the reddish brown spots on the body and the many small reddish papillae of similar size anteriorly and posteriorly. From Baird's description, Selenka, de Man, and Billow suggest in their Appendix E that Phascolosoma fasciatum Baird may be the same as Phascolosoma granulatum (Leuckart) 1828. DESCRIPTION : In both syntypes the introvert is partially retracted ; the trunk of one specimen measures 30 mm in length with a maximum width of 5 mm and that of the other is 20 mm long with a maximum width of approximately 4 mm. The exposed portion of the introverts of the two specimens are marked by dorsal reddish brown bands and pale, inconspicuous papillae, becoming larger and more numerous toward the base of the introvert. In the anal region the papillae are variable in size and shape, the larger ones being pyrimidal and the others flat and low. The papillae of the middle trunk are smaller and rounded, whereas those of the posterior trunk are comparable in size and shape to the anal papillae. The smaller syntype was dissected (by ACS) and an incision was made in the anter- ior introvert for a study of the hooks and small papillae lying between the hooks. The hooks, measuring approximately 0-072 mm at the base and 0-074 mm high, are SIPUNCULA AND ECHIURA 61 characterized by a well-developed central clear streak which shows no basal expansion and a marked terminal curvature with a secondary tooth (Figure 17). The longitudinal muscle bundles in the dissected syntype number 22 immediately anterior to the origin of the dorsal retractors and show little anastomosis. A pair of ventral retractor muscles originates in the posterior third of the trunk and a pair of dorsal retractors in the middle third. The roots of the ventrals span muscle bundles 2 to 8 and the dorsals 5 to 8. A fixing muscle arises left of the ventral nerve cord anteriad to the origin of the dorsal retractors and divides into two branches, one attaching to the rectum and one to the first descending intestinal coil. The nephridia open at the level of the anus and extend posteriorly to the origin of the dorsal retractors. They are attached for approximately a half of their length. REMARKS : The characters as observed in this examination of the syntypes lend support to Selenka's suggestion that Phascolosoma fasciatum Baird 1868 is identical with Phascolosoma granulatum (Leuckart) 1828. The internal anatomy is essentially the same as described by Selenka for P. granulatum and the small papillae of the introvert (Figure 16) and the hooks (Figure 17) correspond in structure to those figured by Selenka for P. granulatum (Selenka, de Man, and Biilow, 1883, PI. x, Figures 147-149). Phascolosoma Jeffrey sii Baird, 1868 Phascolosoma jeffreysii Baird, 1968, p. 88-89. HOLOTYPE : Reg. No. 1863.12.4.8. TYPE LOCALITY : Spezzia. Coll. J. G. Jeffreys. Baird (1868) defined the species on the basis of its shape, the red markings on the dorsal body, transverse striations, density of papillae on posterior and anterior trunk, sparsity of papillae on introvert, and the dorsal reddish brown rings of the introvert. His reasons for distinguishing Phascolosoma jeffreysii as a species separate from P. fasciatum are not clear. The following description includes internal anatomy and is intended to supplement Baird's report. DESCRIPTION : The body length of the holotype is 40 mm with approximately 10 mm of the introvert exposed and the remainder retracted. Papillae at the base of the introvert are pyrimidal in shape and reddish-brown in colour ; at the posterior extremity many are similar, but others are rounded and colourless. Papillae over the rest of the body are mostly smaller, flat, and colourless. Hooks on the retracted introvert possess an accessory tooth and a clear central streak with little basal expansion (Figure 19). They measure approximately 0-054 mm at their base and 0-049 mm m height. The holotype was dissected by one of the present authors (ACS) . The longitudinal musculature is divided into 20 bundles in the region of the origin of the ventral retractors. The ventral retractors originate in the posterior third of the body ; the root of the left ventral retractor spans bundles 2-6 and the right 2-7. The dorsal 62 MARY E. RICE & A. C. STEPHEN retractor muscles originate more anteriorly in the middle third of the trunk and their roots span bundles 4-7. A fixing muscle with two branches attaches to the rectum and to the first descending coil of the gut. The spindle muscle attaches posteriorly to the body wall and anteriorly it attaches immediately in front of the anus. Two nephridia open at the level of the anus and are attached to the body wall for two- thirds of their length. They extend posteriorly to the origin of the ventral retractor muscles. REMARKS : This specimen exhibits striking similarities to the syntypes of Phasco- losoma fasciatum Baird and, like them, to descriptions of P. granulatum (Leuckart) in the shape and distribution of papillae, pigment markings on the trunk and intro- vert, and internal anatomy. The hooks are similar in structure to those of P. fasciatum (Figures 19, 17), except that the size and relative proportions vary and the clear area on the concave side is more distinct. Without seeing the specimen Selenka, de Man, and Biilow (1883) suggested that P. Jeffrey sii Baird 1868 might be the same as P. granulatum (Leuckart) 1828. Phascolosoma lordi Baird, 1868 Phascolosoma lordi Baird, 1868, p. 92-93. HOLOTYPE : Reg. No. 1860.3.21.75. TYPE LOCALITY : Esquimalt Harbour, Vancouver Island. Coll. J. K. Lord. Baird (1868) suggested that this specimen, found in the same locality as two speci- mens which he identified as P. agassizii, might be a variety of the latter. He dis- tinguished P. lordi as a separate species on the basis of differences in " general appear- ance, size, and colour ". In Appendix E of their monograph, Selenka, de Man, and Biilow (1883) stated that P. lordi Baird seemed to be a variety of P. agassizii Kefer- stein. Fisher (1952) in his treatise on " Sipunculid Worms of California and Baja California " gave an exhaustive account of P. agassizii and included P. lordi in the synonymy. Internal anatomy of Baird's holotype, not heretofore reported, is similar to Phasco- losoma agassizii. Longitudinal muscle bundles exhibit a high degree of anastomosis ; 20 bundles were counted between the origins of the ventral and dorsal retractor muscles. The ventral and dorsal retractors originate in the posterior third of the trunk, the dorsals slightly anterior to the ventrals, and the muscles on each side soon fuse to form two muscles which continue separately to their union in the anterior introvert. A spindle muscle attaches to the posterior extremity of the trunk and anteriorly it adheres along the length of the rectum, attaching to the body wall immediately anterior to the anus. One fixing muscle is present, but accurate observations of its attachments were precluded by the fragile condition of the gut. A hook from the anterior introvert of the holotype is illustrated (Figure 18). A comparison with Fisher's (1952) illustrations of hooks of P. agassizii shows similarities to most of the latter in the lack of a secondary tooth, the course and relative width of the central clear streak, and the basal triangular clear streak on the convex side. SIPUNCULA AND ECHIURA 63 Phascolosoma nigriceps Baird, 1868 Phascolosoma nigriceps Baird, 1868, p. 90, pi. xi, figs, i, la. Phymosoma antillarum : Selenka, de Man, and Billow, 1883, p. 58. SYNTYPES : Reg. No. 1859.12.7.63 a/b. TYPE LOCALITY : St. Thomas, West Indies. Coll. Cuming (?). Baird lists specimens from St. Thomas, Jamaica, and Chile. Two specimens from St. Thomas are now present in the collections at the British Museum as syntypes and neither of these had been dissected. Selenka, de Man, and Billow report that they examined Baird' s original specimen from Chile, and it seems that this was not re- turned. These authors referred Baird's specimen to Phascolosoma antillarum Grube and Oersted 1858. The two syntypes from St. Thomas, both with retracted introverts, measure 25 mm in length with a maximum width of 7 mm and 45 mm long with a maximum width of 10 mm. Prominent papillae, typically low and flattened with dark brown platelets of similar size, cover the trunk and are largest and most numerous in the anal region. In the larger specimen the posterior papillae are similar to those of the anal region, whereas in the smaller specimen the posterior cuticle is white and thin with only a few small papillae. In both specimens the platelets are scattered over the cuticle among the papillae. The smaller specimen was dissected (by MER). The longitudinal musculature is divided into anastomosing bundles which number approximately 25 immediately posterior to the origin of the retractor muscles. The 4 retractor muscles originate at nearly the same level at the beginning of the posterior third of the trunk. A fixing muscle attaches on the rectum, anterior to a prominent caecum. The rectum is long and the intestine is comprised of approximately 22 single coils. A well-developed contractile vessel with numerous branched villi extends along the oesophagus into the first intestinal coil. The nephridia are attached to the body wall except for the posterior extremity and reach posteriorly to a level slightly below the origin of the retractor muscles. The characters reported above correspond to those of Phascolosoma antillarum Grube and Oersted 1858 and thus give supporting evidence for Selenka's synonymy of P. nigriceps Baird 1868. Phascolosoma perlucens Baird, 1868 Phascolosoma perlucens Baird, 1868, p. 90-91, pi. x, figs. 2, aa Phymosoma varians : Selenka, de Man and Biilow, 1883, p. 70 SYNTYPES : Reg. No. 1847.12.30.11. TYPE LOCALITY : Jamaica, from holes in coral rocks. Coll. Grosse. Of the three extant syntypes, one was figured by Baird in two illustrations of the entire animal showing external form and size. Selenka, de Man, and Biilow (1883, p. 70) list Phascolosoma perlucens Baird in a synonymy of Phascolosoma varians 64 MARY E. RICE & A. C. STEPHEN Keferstein, with the explanatory statement " Die von uns vorgenommene Unter- suchung des BAIRD' schen Originalexamplars ergab, dass diese Art mit dem Ph. varians KEFERSTEIN identisch ist! ". It is improbable that these authors examined any of the extant syntypes of P. perlucens since these specimens do not correspond to P. varians, but rather to P. dentigerum Selenka and de Man (see below). The occasion for the error remains unexplained. DESCRIPTION : The largest of the three specimens, the one figured by Baird, measures 35 mm in length to the base of the introvert, which is almost entirely retracted, and 3-5 mm in maximum width. In the smallest specimen the trunk is 21 mm long with a maximum width of 1-5 mm and the introvert is partially extended to a length of 7 mm. The trunk of the third specimen is 25 mm in length, 2 mm in maximum width, and the partially extended introvert is 10 mm long. The trunk regions of all specimens are pale and whitish with a thin integument through which longitudinal muscle bundles are visible. The preanal regions and the base of the dorsal introvert are markedly darker due to a concentration of reddish brown, conical, sometimes sharply pointed papillae which become progressively smaller and lighter anteriorly. On the ventral introvert the papillae are generally less prominent . Some papillae on the posterior introvert are pointed in a posterior direction. On the anterior half of the trunk, posterior to the anus, the papillae are colourless, low, mostly oval in shape, and widely spaced. The papillae on the posterior half of the trunk are more conical and light brown in colour, increasing in height and density pos- teriorly and becoming sharply pointed at the posterior extremity. Platelets sur- rounding the central opening of the papillae are darker than peripheral platelets. Hooks from the anterior retracted introvert of the largest specimen measure 0-06 1 mm at the base and 0-065 mm m height (Figure 15). They are sharply bent terminally and show an accessory tooth. The clear triangular area is well-defined. One of the specimens, intermediate in size, was dissected (by MER). The longi- tudinal muscle bands show little anastomosis and number 20-22 posterior to the origin of the ventral retractors. The spindle muscle attaches immediately anterior to the anus and is attached to the posterior extremity. There are approximately 16 single intestinal coils. A single fixing muscle originates to the left of the ventral nerve cord, anterior to the roots of the dorsal retractor muscle and gives off a branch to the postcesophageal intestine and one to the rectum. Two nephridia, opening at the level of the anus, extend posteriorly one-half the length of the trunk and attach to the body wall for three quarters of their length. A pair of black eye-spots is present on the brain. Sixteen rows of hooks were counted through the wall of the retracted introvert. REMARKS : Phascolosoma perlucens Baird 1868 corresponds to Phascolosoma denti- gerum Selenka and de Man 1883. Similarities are apparent in the following taxo- nomic characters : form, distribution, and colouration of papillae, structure of hooks (Figure 15) and hook papillae (Figure 14), relative proportions of the body, and essential features of internal anatomy. SIPUNCULA AND ECHIURA 65 Phascolosoma placostegi Baird, 1868 Phascolosoma placostegi Baird, 1868, p. 89-90. HOLOTYPE : Reg. No. 1965.25.11 TYPE LOCALITY : Cape of Good Hope. Coll. Krauss ; found lodged in a mass of Serpulidae (Placostegus) . The holotype, hardened and brittle, is in very poor condition and disintegrates when manipulated. None of the internal characters could be distinguished, nor any hooks recovered. The reason for the poor state of preservation is found in Baird's state- ment that the specimen when found " was dry, but afterwards moistened and put into spirits " (Baird, 1868, p. 90). Since Baird's description was limited to non- specific external characters, the species cannot be defined and the name Phascolosoma placostegi therefore must be considered as a nomen dubium. Phascolosoma planispinosum Baird, 1868 Phascolosoma planispinosum Baird, 1868, p. 93. Phymosoma nigrescens : Selenka, de Man, and Billow, 1883, p. 73. HOLOTYPE : Reg. No. 1965.25.6. TYPE LOCALITY : Unknown. Coll. Cuming. The holotype has been allowed to dry out completely at some time and is quite contracted and inflexible. It had been dissected previously and most of the internal organs are missing in part or entirely. Fortunately the specimen was seen by Selenka, de Man and Biilow (1883) who placed it in synonymy with Phascolosoma nigrescens Keferstein 1865 and considered it to be identical with a variety from the Philippine Islands which they described but to which they did not give a varietal name. Since Baird stated that he had only one specimen, this is undoubtedly the same specimen which Selenka examined. In spite of the poor condition of the specimen, it was possible to recover some hooks, one of which is figured (Figure 20). This hook differs from those of speci- mens of Phascolosoma nigrescens from the Fiji Islands and Mauritius illustrated by Selenka (Selenka, de Man, and Biilow 1883, Figures 130, 135), in that the clear streak of P. planispinosum is quite distinct from the clear triangular area and an accessory tooth is lacking. There is, however, a thickening in the basal plate, similar to that characteristic of P. nigrescens (Fisher 1952). The triangular space was not as distinct in all of the hooks examined as in the one illustrated (Figure 20), but the form of the central clear streak appeared consistent. Selenka, de Man and Biilow did not illustrate hooks from the specimens of P. nigrescens from the Philippines with which they considered P. planispinosum identical. B* 66 MARY E. RICE & A. C. STEPHEN Themiste lageniformis Baird, 1868 Themiste lageniformis Baird, 1868, pp. 98-99, pi. 10, figs. 3-30. SYNTYPES : 1965.25.9/10 TYPE LOCALITY : Australia? The two specimens on which Baird based his original description of the species are in the same condition as he indicated at that time. In one specimen the introvert and tentacles are extended, but the animal is hard and brittle, apparently having been desiccated at some time after preservation. In the second specimen the intro- vert is completely retracted. The second specimen was dissected by one of the pres- ent authors (ACS). Baird noted the resemblance of the shape of the extended body to a flask and from this character he derived the specific name, lageniformis, meaning flask-shaped. Other characters used by Baird to define the species were as follows : striations and folds of the skin approaching a clathrate pattern posteriorly ; a long, cylindrical introvert covered by a wrinkled, plicate skin ; six pinnate tentacles (the number 6 was followed by a question mark). The following description includes information on the internal anatomy of one of the syntypes and is intended to supplement Baird's reported observations. DESCRIPTION : The extended specimen, previously desiccated and grotesquely contracted in the mid-region of the trunk, measures approximately 22 mm in length to the base of the tentacles. The number of tentacles appears to be six, but the specimen was preserved in such a way that the oral disc was not visible and manipula- tion was precluded by the brittle condition ; hence, it was not possible to affirm whether these were six primary stems. Hooks do not occur on the introvert. The anal opening occurs along the narrowed anterior portion of the body, 8 mm from the base of the tentacles, and the introvert is approximately one-third to one quarter the length of the remainder of the body, an exact figure being difficult to ascertain because of the contracted region in the middle of the body. In the other syntype, in which the anterior end is retracted, the length of the trunk from anus to posterior extremity is approximately 24 mm and the withdrawn introvert is about a quarter of this length. Longitudinal and transverse grooves in the skin form a tessellated pattern over most of the trunk. The pattern is most pro- nounced at the base of the introvert and posterior extremity where the grooves are deepest. In the middle of the body the longitudinal grooves are weak or non-existent and the transverse grooves shallow ; consequently, the tessellation is not so apparent The openings of the epidermal glands occur within the rectangles of the tessellation ; in the middle of the body where the skin is more distended they are most readily observed and appear as openings in the centre of concave depressions (Figure 13). Internally the body wall musculature of the dissected syntype is smooth, two thick retractors originate in the posterior fifth of the trunk remaining separate for most of their length, and the gonad appears as a thin strand on the base of the re- tractors (Figure 12). The oesophagus is directed posteriorly to a point at the base of the ventral retractors where it is attached by a fixing muscle ; it then turns SIPUNCULA AND ECHIURA 67 abruptly anteriorly to enter the intestinal coil. The intestine is comprised of approximately 28 single coils and has no posterior attachment. Through the thin walls of the descending gut the contents can be seen to consist of fine sand or mud particles which have been compacted and twisted into a spiral form. A caecum is present at the beginning of the rectum. The spindle muscle, passing by and attaching to the base of the caecum, runs along the dorsal side of the rectum and appears to insert on it. There is no attachment of the spindle muscle to the posterior body wall. Prominent wing muscles fasten the rectum to the body wall in the region of the anus. There are three fixing muscles. Fj attaches the oesophagus to the body wall just posteriorly to the inner margin of the left retractor muscle. Broken from its site of origin on the body wall, F2 is attached to the first descending intestinal coil. F3, attached to the body wall near the dorsal midline in the anterior third of the body, proceeds beneath the intestinal coil to attach to the beginning of the rectum as it emerges from the intestinal coil. A prominent contractile vessel is adjoined to the descending oesophagus but ends just beyond the point at which the oesophagus curves anteriorly toward the intestinal coil. Anteriorly the vessel is very much enlarged and distended with cellular elements, but more posteriorly it gives off numerous tufts of short filiform villi which frequently bifurcate near their basal attachments. Two nephridia hang freely in the body cavity. The nephridiopores open at about the same level as the anus. REMARKS : These specimens correspond to Selenka's (1883) description and illustrations of Dendrostoma signifer in the characteristic arrangement and form of the contractile vessel and villi, the number and attachment of the fixing muscles, the relatively short introvert, and the furrowing of the skin. Although Selenka does not mention a caecum for D. signifer, other authors (Edmonds 1956, Ikeda 1904, Wesenberg-Lund 1959) have reported a rectal diverticulum for this species. The number of tentacular stems has been variously reported as 5 or 6 (Selenka 1883) or as 4 (Ikeda 1904, Edmonds 1956, Fischer 1919). Although Selenka lists Themiste lageniformis in an appendix (Selenka, de Man, and Billow, 1883, Appendix E), he makes no mention of the name elsewhere in the monograph. Presumably he had not examined Baird's syntypes when he described Dendrostoma signifer as a new species. Aspidosiphon cumingii Baird, 1868 Aspidosiphon cumingii Baird, 1868, p. 102, pi. xi, fig. 2 ; Selenka, de Man and Billow, 1883, p. US-US- TYPE LOCALITY : Philippine Islands. Coll. Cuming. Baird's description was superficial, but the holotype was fully described by Selenka, de Man and Billow and the name remains valid. The specimen is missing from the collection, possibly never returned to the British Museum (Natural History) by Selenka. 68 MARY E. RICE & A. C. STEPHEN Aspidosiphon jukesii Baird, 1873 A spidosiphon jukesii Baird, 1873, p. 97. HOLOTYPE : Reg. No. 1965.25.3. TYPE LOCALITY : Lee Sandbanks (Great Barrier Reef, Australia?). Baird's brief description (Baird, 1873) enumerated a few general external features, but provided no significant characters by which this species could be distinguished by other authors. Hence the species was not recognized in the literature, even though the type remained extant in reasonably good condition in the collections of the British Museum (Natural History). The holotype was dissected by one of the present authors (ACS) and a report of its internal anatomy is presented here. The specimen had been removed at the time of its collection from a solitary coral, the remains of which are still preserved. DESCRIPTION : Preserved in a curved, U-shaped position, the specimen measures 25 mm along the median line of the outer, dorsal curvature with a maximum width of approximately 5 mm (Figure 24). The introvert is entirely retracted. The integument of the posterior two-thirds of the trunk is thin, distended, and pale yellow, whereas the anterior one-third is more tightly contracted and a deeper yellow. A well-developed, dorsally oblique anal shield is clearly set off from the remainder of the body (Figure 21, 22). Approximately 12 major longitudinal furrows, not all complete, mark the flattened surface of the shield which is composed of large, irregularly shaped, amber-coloured platelets. Lateral and ventral extensions of the shield are distinguished by raised papillae, densely packed, with embedded amber platelets similar to those of the dorsal surface of the shield. A small clear spot marks the apex of each papilla. Over the remainder of the trunk the papillae are more widely dispersed, smaller, and flatter with a relatively large central area surrounded by one or more rings of pale yellow coalesced platelets which form clumps of varying sizes (Figures 28, 29). The papillae may reach a width of 0-7 mm in the posterior quarter of the trunk, but they are only slightly elevated from the surface with a maximum height of approximately 0-3 mm. The posterior shield is circular, well-demarcated, but lighter in color than the anal shield (Figure 23). There are a few weak, irregular radial striations which do not extend to the center of the shield. The constituent pale yellow platelets vary in size and are larger and darker in the central portion. Small, pale hooks are arranged in rows on the anterior introvert. At their base the hooks measure about 0-028 mm and their height is 0-025 mm. The hooks are weakly curved with a secondary terminal point and a large central clear area which is widened proximally to include the entire base (Figure 25). Introvert papillae, measuring about 0-013 mm in height and 0-008 mm in diameter, are dispersed among the rows of hooks (Figure 27). Larger spines with a height of 0-035 mm are scattered more posteriorly over the introvert. Because of its retracted condition, the entire introvert was not examined. The musculature of the body wall is smooth. Two retractors attach at the posterior extremity in the region of the terminal shield. Separated posteriorly, they soon join and are united for about two-thirds of their total length. At the point of union SIPUNCULA AND ECHIURA 69 of the retractors the oesophagus bends in an anterior direction to join the intestinal coil. Numerous coils (approximately 30 single coils) comprise the intestinal spiral, but an accurate count is not possible because of the poor state of preservation of the gut. A spindle muscle attaches the intestine posteriorly. The anal opening is immediately posterior to the flattened anterior shield and the two nephridia open at nearly the same level. The nephridia are partially attached to the body wall, but unfortunately the free ends have been broken off so that neither the length of the nephridia nor the relative extent of their attached portion can be determined. REMARKS : This specimen resembles Aspidosiphon coratticola Sluiter, 1902, both in its habitat in a solitary coral and in the following taxonomic characters : origin and union of the two retractor muscles, morphology of anterior and posterior shields, and the structure and distribution of the hooks, spines, and papillae. Although similar in basic structure, the hooks and spines differ in size for the holo- type of Aspidosiphon jukesii and A . coratticola. The height of the hooks is the same in both (0-025 mm), but the width in A. corallicola is proportionately greater. Moreover, Sluiter (1902) reported that in A. corallicola the spines were smaller than the hooks, whereas the reverse is true for the holotype of A. jukesii. Without additional measurements to indicate the range of variation within and among indivi- duals the significance of the size discrepancies is difficult to evaluate. However, the other important similarities between A . jukesii and A . corallicola suggest that the two names represent a single species. A . jukesii has priority, since it is the older name. Pseudasipidosiphon gracile Baird, 1868 Pseudasipido siphon gracile Baird, 1868, p. 103, pi. x, fig. i, la. Aspidosiphon gracilis : Selenka, de Man and Billow, 1883, p. 122-123, pi. ii, fig. 22, pi. xiv, fig. 209-213. SYNTYPES : Reg. No. 43.5.i5.58a/b. TYPE LOCALITY : Philippine Islands. Coll. Cuming. Baird seems to have had three specimens, one of which was examined and described by Selenka. Since neither of the two specimens in the collection has been previously dissected, it is probable that Selenka did not return the specimen which he described. Selenka gave a complete, well-illustrated description of Baird's specimen, so that the specific name remains valid. Echiurus farcimen Baird, 1873 Echiurus farcimen Baird, 1873, p. 97. Echiurus chilensis : Shipley, 1899, p. 342. HOLOTYPE : Reg. No. 69.6.28.18. TYPE LOCALITY : Punta Arenas, Patagonia. Coll. Cunningham. In Baird's brief description of the species, he mentioned 5 specimens, the largest of which was 16 inches in length. Only one of these specimens remains in the collection and, although previously dissected, it is still in good condition. Its approximate measurements are 170 mm in length and 125 mm in maximum circumference. With yo MARY E. RICE & A. C. STEPHEN rounded extremities and considerably reduced prostomium, the specimen resembles a sausage in shape. One of the members of the anteroventral pair of setae is missing, but the intact seta, slightly curved, is extended to a length of 4 mm. A single ring of 12 smaller setae, marked by a mid-ventral gap, encircles the posterior extremity. There are three pairs of prominent nephridia, each with two long, spirally coiled lips. Two anal vesicles reach lengths approximately one-half that of the body. Shipley (1899) considered Baird's specimens of Echiurus farcimen to be synony- mous with E. chilensis Max Miiller 1852 and later authors (Fisher 1946, Wesenberg- Lund 1955, Amor 1965) have accepted this synonymy. Since 1907, when Seitz revised the genus, this species has been known as Urechis chilensis.1 TABLE I 1. Gray and Baird species-names which are senior subjective synonyms of currently accepted names. Senior synonym Current name Siphunculus arcuatus Gray, 1828 Phascolosoma lurco (Selenka & de Man, 1883) Phascolosoma perlucens Baird, 1868 Phascolosoma dentigerum (Selenka & de Man, 1883) Themiste lageniformis Baird, 1868 Themiste signifer (Selenka & de Man, 1883) Aspidosiphon jukesii Baird, 1873 Aspidosiphon corallicola Sluiter, 1902 2. Gray and Baird species-names which are currently accepted. Themiste hennahi Gray, 1828 Sipunculus angasi Baird, 1868 Phascolosoma albolineatum Baird, 1868 Aspidosiphon cumingi Baird, 1868 Aspidosiphon gracile (Baird, 1868) 3. Gray and Baird species-names which are junior subjective synonyms of currently accepted names. Junior synonym Current name Siphunculus dentalii Gray, 1828 Phascolion strombi (Montagu, 1804) Siphunculus tuberculatus Gray, 1828 Phascolosoma noduliferum Stimpson, 1855 Sipunculus aeneus Baird, 1868 Siphonosoma australe (Keferstein, 1865) Sipunculus deformis Baird, 1868 Siphonosoma cumanense (Keferstein, 1866) Sipunculus eximioclathratus Baird, 1868 Sipunculus nudus Linnaeus, 1766 Phascolosoma aethiops Baird, 1868 Phascolosoma antillarum Grube & Oested, i. Seitz (1907) erected the genus Urechis for the species Echiurus chilensis Max Miiller and E. unicinctus von Drasch. Riveras Zuniga (1942), on grounds of priority, revived the generic name Pinuca Hup6 in Gay 1854, but more recently Jones, Hedgpeth, and Hand (1968) have applied to the International Com- mission on Zoological Nomenclature for the suppression of Pinuca. At this date no action has been taken on this request. SIPUNCULA AND ECHIURA 71 Junior synonym Current name Phascolosoma capsiforme Baird, 1868 Golfingia margaritacea (Sars, 1851) Phascolosoma fasciatum Baird, 1868 Phascolosoma granulatum (F. S. Leuckart, 1828) Phascolosoma grayi Baird, 1868 (nom. Phascolosoma noduliferum Stimpson, 1855 nov. pro. Siphunculus tuberculatus Gray, 1828) Phascolosoma jeffreysii Baird, 1868 Phascolosoma granulatum (F. S. Leuckart, 1828) Phascolosoma lordi Baird, 1868 Phascolosoma agassizii Keferstein, 1866 Phascolosoma nigriceps Baird, 1868 Phascolosoma antillarum Grube & Oersted, 1859 Phascolosoma planispinosum Baird, 1868 Phascolosoma nigrescens Keferstein, 1865 Echiurus farcimen Baird, 1868 Urechis chilensis Max Miiller, 1852 4. Baird name which is a nomen dubium. Phascolosoma placostegi Baird, 1868. REFERENCES AMOR, A. 1965. Una neuva localidad para Pinuca chilensis (Max Mtiller) en el Atlantico Sur (Echiurida). Aclaracidn sobre su sinonima : Pinucidae nom. nov. para Urechidae Fisher and MacGinitie. Physis, B. Aires, 25 : 165-168. (In Press). A prop6sito del hellazgo de Themiste hennahi Gray en la Bahfa Concepcidn, Chile (SIPUNCULA). BAIRD, W. B. 1868. Monograph of the species of worms belonging to the sub-class Gephyrea ; with a notice of such species as are contained in the collection of the British Museum. Proc. zool. Soc. Lond. 1868 : 76-114. — 1873. Description of some new species of Annelida and Gephyrea in the collection of the British Museum. /. Linn. Soc. Zool. 11 : 94-97. BENHAM, W. B. 1903. The sipunculids of New Zealand. Trans. Proc. N.Z. Inst. 36 : 172-184. 1922. Gephyrean inermia. Australian Antarctic Expedition 191 1-1914, under the leader- ship of Sir Douglas Mawson. Scient. Rep. Australas. Antarct. Exped. Ser. 6. Zool. and Bot. 6 : 1-22. COLLIN, A. 1892. Gephyreen gesamelt von Herrn aber-Stabsarzt Dr. Sander auf der Reise S.M.S. " Prinz Adalbert ". Arch. Naturgesch, 1 : 177-182. EDMONDS, S. J. 1955- Australian Sipunculoidea. i. The genera Sipunculus, Xenosiphon, and Siphonosoma. Aust. J. mar. Freshwat. Res. 6 : 82-97. 1956. Australian Sipunculoidea. 2. The genera Phascolosoma, Dendrostomum, Golfingia, Aspidosiphon, and Cleosiphon. Aust. J. mar. Freshwat. Res. 7 : 281-315. 1961. On Sipunculus aeneus Baird (Sipunculoidea). Ann. Mag. not. Hist. Ser. 13, 4 : 217-220. 1965. Sipunculoidea of the Ross Sea. N.Z. Oceanographic Institute Memoir No. 27. Bull. N.Z. Dept. sclent, ind. Res. 167 : 27-33. FISCHER, W. 1896. Gephyreen. (4) 1-7 in Ergebnisse der Hamburger Magalhaensische Sam- melreise. 1 : Hamburg, Friederichsen. 1913. Ueber einige sipunculiden des naturhistorischen museums zu Hamburg. Jb. hamb. wiss. Anst. 30 : 93-101. 1914- Weitere Mitteilungen iiber die Gephyreen des Naturhistorischen (Zoologischen) Museums zu Hamburg. Mitt. zool. Mus. Hamb. ser. 2, 31 : 1-28. Gephyreen der Siidwestkiiste Australiens. Zool. Anz. 50 : 277-285. 72 MARY E. RICE & A. C. STEPHEN FISHER, W. K. 1946. Echiuroid worms of the North Pacific Ocean. Proc. U.S. nat. Mus. 96 : 215-292. — 1952. The sipunculid worms of California and Baja California. Proc. U.S. natn. Mus. 102 : 371-450. GEROULD, J. H. 1913. The sipunculids of the eastern coast of North America. Proc. U.S. natn. Mus. 44 : 373-437. GRAY, J. E. 1828. Spicilegia Zoologica ; or original figures and short systematic descriptions of new and unfigured animals, (i) 1-8. London : Treiittel, Wiirtz and Co. ; and W. Wood. IKEDA, I. 1904. The gephyrea of Japan. /. Coll. Sci. imp. Univ. Tokyo, 20(4) : 1-87. JOHNSTON, G. 1833. Illustration of British Zoology. Mag. nat. Hist., 6 : 232-235. JONES, M. L., HEDGPETH, J. and HAND, C. 1968. Pinuca Hupe in Gay, 1854 (Echiuroidea) : Proposed suppression under the plenary powers. Z. N. (S.) 1836. Bull zool. Nom. 25 : 100-102. LANCHESTER, W. F. 1905. On the sipunculids and echiurids collected during the " Skeat " expedition to the Malay Peninsula. Proc. zool. Soc. Lond. 1 : 35-41. RIVEROS ZUNIGA, F. Pinuca chilensis. Prensas Univ. Chile. 1-15. SEITZ, P. 1907. Der Bau von Echiurus chilensis (Urechis n. g. chilensis). Zool. Jb. Anat. 24 : 323-356. SELENKA, E., DE MAN, J. G. and BULOW, C. 1883. Die Sipunculiden. Zweiter Theil. Wissenschaftliche Resultate. 4 : (i) 1-131. in Semper, C. G. Reisen im Archipel der Philippinen. Leipzig & Weisbaden. SHIPLEY, A. E. 1899. On a collection of echiurids from the Loyalty Islands, New Britain, and China Straits, with an attempt to revise the group and to determine its geographic range. In Willey, A. Zoological results based on material from New Britain, New Guinea, Loyalty Islands and elsewhere collected during the years 1893, i8g6 and 1897. 3 : 335-356. London : Cambridge University Press. SLUITER, G. P. 1902. Die Sipunculiden and Echiuriden der Siboga-Expedition nebst zusam- menstellung der ueberdies aus den indischen Archipel bekannten Arten. Siboga-Exp. 25 : 1-53. STEPHEN, A. C. 1964. A revision of the classification of the phylum Sipuncula. Ann. Mag. nat. Hist. Ser. 13, 7 : 457-462. TEN BROEKE, A. 1929. Sipunculoidea und Echiuroidea. Tierwelt Dtl. 15 : 156-168. WESENBERG-LUND, E. 1955. Gephyrea from Chile. Reports of the Lund University Chile Expedition of 1948/49, 19. A eta Univ. lund. N.F. 51 : (10) 1-24. 1959. Campagne 1956 du " Calypso " dans le Golfe de Guinee et aux lies Principes Sao Tome et Annobon. Sipunculidea and Echiuroidea. Annls. Inst. oceangr. Monaco. 37 : 207-217. MARY E. RICE, Ph.D., Associate Curator, Division of Worms, NATIONAL MUSEUM OF NATURAL HISTORY, SMITHSONIAN INSTITUTION, WASHINGTON, D.C. 20560, UNITED STATES OF AMERICA. f Dr A. C. STEPHEN, D.Sc., F.R.S.E., Keeper of Natural History, ROYAL SCOTTISH MUSEUM, CHAMBERS STREET, EDINBURGH, i, SCOTLAND. t Obituary : Waterston, A. R. 1966, Nature, 211 : 21. PLATE i Siphunculus arcuatus Gray. Holotype. Current Name : Phascolosoma arcuatum (Gray) FIG. i. Dissected specimen showing internal anatomy. In the preserved specimen the retracted introvert is coiled within the body cavity ; for clarity in the drawing it is straightened and the intestinal coil is pulled aside to reveal other internal structures. AR, anterior retractor ; E, oesophagus ; G, gonad ; I, retracted introvert ; N, nephridium ; NC, ventral nerve cord ; PR, posterior retractor ; R, rectum ; S, spindle muscle ; W, wing muscle. Approximate length of specimen is 100 mm. FIG. 2. Papilla from middle region of trunk. Note polygonal platelets, smaller and more concentrated around centre, but otherwise evenly distributed. Diameter 0-3 mm. FIG. 3. Lateral view of holotype, showing external features. Introvert is retracted. A, anus ; NP, nephridiopore. Rectangles a, b, and c, each measuring 1x4 mm on specimen, are enlarged at left to show relative size, form and distribution of papillae. 3a. Enlargement of skin area at base of introvert. 3b. Enlargement of skin area from middle region of trunk. 3C. Enlargement of skin area from posterior extremity. FIG. 4. Diagrammatic representation of body wall from middle third of trunk, showing the coelomic sacs and their relation to the musculature of the body wall and overlying integument. The integument, shown only in the lower right portion of the diagram, is markedly thinner in the areas covering the sacs. CM, circular muscle bundle ; CS, coelomic sac ; IN, integument ; LM, longitudinal muscle bundle ; P, papilla. FIG. 5. Hook from introvert. 0-08 x 0-07 mm. Bull. Br. Mus. nat. Hist. (Zoot.) 20, 2 PLATE i m PLATE 2 FIG. 6. Themiste hennahi Gray. Paralectotype, dissected specimen. A, anus ; C, caecum ; CV, villi of contractile vessel ; E, oesophagus (looped under right retractor) ; Fi, fixing muscle i ; Fa, fixing muscle 2 (broken) ; F3, fixing muscle 3 ; N, nephridium ; NC, nerve cord ; RM, retractor muscle ; S, spindle muscle. Approximate length of trunk is 37 mm. FIG. 7. Themiste hennahi Gray. Lectotype. Oral view of tentacular crown, oriented with dorsal tentacles toward bottom of page. The 6 lips surrounding the mouth are evident between the bases of the tentacles. FIG. 8. Themiste hennahi Gray. Lectotype. Lateral view showing external features. A, anus ; NP, nephridiopore. Approximate length of body (minus tentacles) is 70 mm. Squares a, b, c, d, each i mm2 on specimen, are enlarged on right to show details of skin and papillae. 8a, Introvert. Cuticle has been detached exposing canals of epidermal papillae. 8b, Enlarge- ment of integument in anal region. 8c, Enlargement of integument in middle of trunk. 8d, Enlargement of integument in posterior extremity of trunk. FIG. 9. Siphunculus tuberculatus Gray. Holotype. Current name : Phascolosoma noduli- ferum Stimpson. Hook from introvert. 0-058 mm x 0-047 mm. FIG. 10. Siphunculus tuberculatus Gray. Holotype. Papilla from base of introvert. Diameter, 0-27 mm. Darkly pigmented platelets are arranged in a distinctive ring around the clear central area. FIG. ii. Siphunculus tuberculatus Gray. Holotype. Posterior i/io of trunk, showing dense concentration of prominent, dome-shaped papillae. Bull. BY. Mus. nat. Hist. (Zool.) 20, 2 PLATE 2 PLATE 3 FIG. 12. Themiste lageniformis Baird. Syntype. Dissected specimen. In the specimen the incision had been made through the outer edge of the base of the left retractor muscle, but for purposes of clarity and orientation in the drawing the line of incision and the muscle are reconstructed so that the muscle appears intact. C, caecum ; CV, contractile vessel ; E, oesophagus ; Fi, fixing muscle i ; F2, fixing muscle 2 (broken) ; F3, fixing muscle 3 ; G, gonad ; N, nephridium ; NC, nerve cord ; R, rectum ; RM, retractor muscle. Approximate length of trunk is 24 mm. FIG. 13. Themiste lageniformis Baird. Syntype. Enlargement of skin (i mm2) from pos- terior area of trunk showing horizontal grooves and less distinct vertical grooves surrounding openings of epidermal glands. FIG. 14. Phascolosoma perlucens Baird. Syntype. Apical view of small papilla from introvert, located among rows of hooks on anterior introvert. FIG. 15. Phascolosoma perlucens Baird . Syntype. Hook from anterior introvert. 0-061 x 0-065 mm. FIG. 16. Phascolosoma fasciatum Baird. Syntype. Current name : P. granulatum (Leuck- art). Lateral view of small papilla from introvert ; situated among rows of hooks on anterior introvert. FIG. 17. Phascolosoma fasciatum Baird. Syntype. Current name : P. granulatum (Leuck- art). Hook from introvert. 0-072 mm (base) x 0-074 mm (height). FIG. 18. Phascolosoma lordi Baird. Holotype. Current Name : P. agassizii Keferstein. Hook from introvert. 0-057 x o-o68 mm. FIG. 19. Phascolosoma jeffreysii Baird. Holotype. Current name : P. granulatum (Leuck- art). Hook from introvert. 0-057 X 0-049 mm. FIG. 20. Phascolosoma planispinosum Baird. Holotype. Current Name : P. nigrescens Keferstein. Hook from introvert. 0-053 x 0.041 mm. FIG. 21. Aspidosiphon jukesii Baird. Holotype. Laterodorsal view of anterior shield. A, anus. FIG. 22. Aspidosiphon jukesii Baird. Holotype. Lateral view of anterior shield. A, anus. FIG. 23. Aspidosiphon jukesii 'Baird. Holotype. Apical view of posterior shield. FIG. 24. Apsidosiphon jukesii Baird. Holotype. Lateral view of entire animal. Specimen is approximately 25 mm in length, measured along median dorsal curvature. FIG. 25. Aspidosiphon jukesii Baird. Holotype. Hook from anterior introvert. 0-028 (base) x 0-025 mm (height). FIG. 26. Aspidosiphon jukesii Baird. Holotype. Spine from posterior introvert. 0-035 mm (height). FIG. 27. Aspidosiphon jukesii Baird. Holotype. Lateral view of small papilla from introvert ; situated among rows of hooks on anterior introvert. FIGS. 28, 29. Aspidosiphon jukesii Baird. Papillae from the middle region of the trunk showing coalescence of platelets. 28. 0-54 mm, diameter. 29. 0-81x0-54 mm. Bull. Br. Mus. nat. Hist, (Zool.) 20, 2 PLATE 3 Printed in Great Britain by Alden & Mowbray Ltd at the Alden Press, Oxford THE TYPES AND FIGURED SPECIMENS OF UNIONACEA (MOLLUSCA : BIVALVIA) IN THE BRITISH MUSEUM (NATURAL HISTORY) R. I. JOHNSON BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 20 No. 3 LONDON : 1971 THE TYPES AND FIGURED SPECIMENS OF UNIONACEA (MOLLUSCA : BIVALVIA) IN THE BRITISH MUSEUM (NATURAL HISTORY) BY RICHARD IRWIN JOHNSON J ---- Museum of Comparative Zoology, Harvard University Pp. 73-108 ; 2 Plates BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 20 No. 3 LONDON : 1971 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1 949, is issued in five series corresponding to the Departments of the Museum, and an Historical series. Parts will appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. In 1965 a separate supplementary series of longer Papers was instituted, numbered serially for each Department. This paper is Vol. 20 No. 3 of the Zoological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation Bull. Br. Mus. nat. Hist. (Zool.). Trustees of the British Museum (Natural History), 1971 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 20 January, 1971 Price £1.20 THE TYPES AND FIGURED SPECIMENS OF UNIONACEA (MOLLUSCA : BIVALVIA) IN THE BRITISH MUSEUM (NATURAL HISTORY) By RICHARD I. JOHNSON INTRODUCTION AMONG the more important collections of Unionacea in the British Museum (Nat. Hist.) is that of Hugh Cuming, which was acquired in 1866. Cuming sought to have a pair of each described species, and he exchanged shells with most of his contempor- aries. Many of his examples of North American species were from J. G. Anthony, whose collection is in the Museum of Comparative Zoology. These specimens often have generalized locality data, which, with the names, were written on the shells by Anthony. Many of Cuming's shells, and those from the collections of S. Hanley, T. L. Taylor and the Sowerby family, are figured by Lovell Reeve and George B. Sowerby in their uncritical monographs on the Unionacea in the Conchologia Iconica between the years 1864-1870. These authors generally figured shells under the names which appeared on the labels before them. Unnamed ones were apparently described as new species. References to the original descriptions are often scanty or missing, and locality data, when present, usually general. Nevertheless, because of the quality of the hand coloured lithograph plates of the various species in natural size, the work has been often referred to by subsequent authors. The types of Reeve and Sowerby are assumed to have been in the British Museum (Nat. Hist.) if they were based on shells from the collections of Cuming, Hanley or Taylor, though many of these are now missing. The entire Cuming collection was acquired by the museum shortly after his death, but only parts of the collections of Hanley and Taylor were added toward the end of the nineteenth century, after they had passed through the hands of shell dealers. Nevertheless, references to all of the species described from these collections are included in this list if they were described in the Conchologia Iconica. None of the species described by Sowerby in this work from the Sowerby family collection were found in the museum. The Sowerbys were shell dealers and it is assumed that these types were sold to various collectors, and therefore, as they were never in the British Museum (Nat. Hist.), they are not included here. The molluscs described by Alcide D. d'Orbigny in his Voyage dans I'Amerique Meridionale during the years 1826-33, were purchased by the museum and enumer- ated by J. E. Gray in 1854. 1 In most instances Orbigny marked his figured types 1 Gray, John Edward, 1854, List of the Shells of South America in the Collection of the British Museum. Collected and described by M. Alcide d'Orbigny, " In the Voyage dans I'Amerique M£ridonale. " London : Published by Order of the Trustees. Pp. 89, 16 mo. (Unionidae, pp. 74-79, nos. 660-700). 76 R. I. JOHNSON with an " x ". The names of all of the Unionacea described by Orbigny from his collection of this voyage are included, including those for which no types were found. The shells of Orbigny have always been kept separate from the main collection. The specimens were originally glued on cards, but when I examined them, many were detached and mixed ; however, it was possible to sort them. The shells were cleaned, registered and put in boxes with their original labels. Many of Arthur Morelet's types were purchased in 1892, shortly after his death, from the shell dealer Hugh Fulton. Also purchased were those species described from the Morelet collection by Fischer and Crosse in their Mission Scientifique au Mexique et dans I'Amerique Centrale (Unionacea, 1894) . Only those species for which types were found in the museum are included here. While the collections mentioned above contain the majority of types that can be discussed here, there are many types from various individuals that can be found only by reference to the following list. The general collection of Unionacea had been hastily moved during the Second World War, and many of the shells were detached from the cards on which they had been glued and were badly mixed. Fortunately many of the Hanley, Taylor and Morelet shells were in glass topped boxes. Originally most of the lots consisted of two specimens, often accompanied with scanty data. I was unable to rearrange the general collection, but each lot was examined, and every specimen that gave any evidence that it might be a type, or could have been one that was ever figured in the Conchologia Iconica or any other work, was checked against original references, and those which proved to be authentic were placed in a special cabinet which was made available for that purpose. In addition to the list of types, this paper includes references to several works containing figures of specimens which are not types ACKNOWLEDGEMENTS I am grateful to Dr. Norman Tebble who invited me to make the present study, which was done in 1963. He kindly permitted me to remove the types and figured specimens from the general collection, and provided a separate cabinet to house them. He was most kind in assisting me with my many queries, as was Mr. Peter Dance. Miss Joan Rosling painstakingly catalogued and individually numbered all of the shells which were previously unregistered. Mr. Fred Woodward helped me locate some of the figured specimens from the Cuming collection. Thanks are also extended to my colleagues Drs. Kenneth J. Boss and Ruth D. Turner who critically read the manuscript. Special thanks are extended to Mr. John F. Peake for his part in helping to shep- herd this paper through the press, for reading it with special care, and offering critical suggestions. Part i A LIST OF THE TYPES OF RECENT UNIONACEA IN THE BRITISH MUSEUM (NATURAL HISTORY) WITH THEIR ORIGINAL REFERENCES AND TYPE LOCALITIES The following list is arranged alphabetically by species, giving the author, year and place of publication of the original description, the type locality and collector or UNIONACEA: TYPES AND FIGURED SPECIMENS 77 collection when relevent, and includes references to those species which are believed to have been in the museum but are now lost. The word " lost " is in brackets in- dicating this to be my opinion after a careful search had been made. The location of the holotype is given when known, even if not in the British Museum (Nat. Hist). If the location of the holotype is unknown, types are listed as syntypes or paratypes, generally the latter if the species was originally figured and there seemstobea possibility that that specimen may be extant . Obviously the use of either of these terms when the location of the holotype is not definitely known is arbitrary. The selection of a single specimen, or holotype, to represent each described taxon, which is now prevalent, was not necessarily the intent of earlier authors. Never- theless, the rules promulgated by the XV International Congress of Zoology and published as the International Code of Zoological Nomenclature (1961 and slightly modified in 1964) must be uniformly applied to all taxa. Article 73 (a) states : If a new nominal species is based on a single specimen, that specimen is the " holotype ". Clearly covered by this article are most of the Unionacea described and figured by Reeve and Sowerby in the Conchologia Iconica, since the type lot usually consisted only of the figured specimen. There is usually no evidence in the descriptions that they saw any but the figured specimens. Isaac Lea often mentioned the number of specimens that Cuming sent to him, sometimes a single individual. Yet, some lots of specimens were found in the collection which might pass for types, except that the species was based on the single specimen sent to Lea which is now in the United States National Museum. Some " type " lots contained specimens in excess of the number which were mentioned as having been seen by the describer. The authenti- city of paratypes cannot always be ascertained, even in a work such as the present one, but if the holotype has been located their authenticity is less germane. Article 73 (b) states : If an author states in the description of a new nominal species that one specimen and only one is " the type " or uses some equivalent expression, that specimen is the holotype. The key to the sensible application of the concept of a single specimen as the holotype to taxa described from 1756 until this concept was firmly established depends on the spirit in which the phrase " equivalent expression " is interpreted. The semantics of this seemingly ambiguous phrase can be argued, but among the definitions of these words are the following : " equivalent " — alike in significance ; " expression " — act or process of representing or making manifest, especially by language. It was surely not the intent of the Congress to have included this phrase to obfus- cate the recognition of holotypes. Rather, it appears to be a clear mandate to use Occam's razor (entities are not to be multiplied without necessity) in their recog- nition with taxa published before 1961. Thus if a taxon was described with a single set of measurements and not figured, the measured specimen is regarded here, as the holotype. Similarly, if a taxon is described and a single specimen is figured, that specimen is regarded as the holotype. If the measured type cannot be located, a lectotype should be chosen and figured. If the figured type is lost, the selection of a 78 R. I. JOHNSON lectotype can be made, but it may be redundant to do so if the species is readily recognizable from the figure. Article 73 (c) states : If a new nominal species has no holotype under the provisions of (a) and (b) all of the specimens of the type series are " syntypes ", of equal value in nomenclature. Recommendation 746 suggests : A zoologist should choose as lectotype a syntype of which a figure has been published, if such exists. This recommendation can be easily complied with if the author figured more than one specimen that can be located. Occasionally, a subsequent author has figured a type, sometimes without design, in the case of previously unfigured species, which can be selected. It is further recommended (74D) : When possible a lectotype should be chosen from syntypes in the collection of a public institution, preferably of the institution containing the largest number of syntypes of the species, or containing the collection upon which the author of the nominal species worked, or containing the majority of his types. Recommendation 740 presents both ethical and practical problems, but if Article 73 (b) is interpreted as suggested above, as it is in this paper, these problems are minimal. The references to Isaac Lea's Observations on the Genus Unio do not include plates and figures, since they are always the same as the preceding reference. Only the pages were renumbered in this reprint. All locality data, contained in brackets, are additions to already published records and are from original labels or modern atlases. The following abbreviations have been used in this list : ANSP Academy of Natural Sciences of Philadelphia, Pennsylvania. BMNH British Museum (Nat. Hist.). London. MCZ Museum of Comparative Zoology, Cambridge, Massachusetts. USNM Unites States National Museum, Washington, D.C. abdalliana Bloomer, Caelatura aegyptiaca : 1946, Proc. Mai. Soc. London, 27 : 70, pi. 6, figs. 4-5 (Hag Abdalla, north of Sennaar, Soudan). Paratype BMNH 1948.5.5.6. abnormis Morelet, Unio : 1862, Rev. etMag. de Zool. (2) 14 : 480 (Bangkok [Thailand]) ; 1875, Series Conch., pt. 4, p. 347 [not fig.]. Syntype BMNH 93.2.4.1374. acrorrhynchus Martens, Unio : 1895, Sitzber. Gesell. Natur. Freunde, Berlin, p. 214 (Fluss Naemingang bei Hatanggyong [siidlich von Keumsan, Prov. Chollado] und in einem Zufluss des Imjingang [6 km. von Inchon, Prov. Kangwondo] Korea) ; Martens 1905, Zool. Jahrb., Suppl. 8, p. 61, pi. 3, fig. 4. Paratype BMNH 94.11.24.11. acuminatus H. Adams, Unio : 1866, Proc. Zool. Soc. London, p. 376 ([Lake] Albert N'yanza [Central Africa]). Holotype BMNH 1867.1.9.4, figured by Smith, 1892, Ann. and Mag. Nat. Hist. (6) 10, pi. 10, fig. 12, consists of one valve. aegyptiaca Pallary, Mutelina : 1924, M6m. Inst. d'Egypt, 7 : 52, pi. 4, fig. 14 (Canal Mahmou- dieh [Lower Nile, Egypt]). 2 Paratypes BMNH 1937.12.30. I3I55-56- aequatorius Morelet, Unio : 1885, Jour, de Conch., 33 : 31, pi. 2, fig. 9 (la riviere Mayumba, district de Cacongo [Congo] a 3 degree au-dessus de 1'Equateur). Holotype BMNH 93.2.4. 1585 ; paratype BMNH 93.2.4.1586. aereus Reeve, Unio : 1865, Conch. Iconica, 16, Unio, pi. 30, species 160 (Hab.?. Cuming colln.). BMNH [lost]. UNIONACEA: TYPES AND FIGURED SPECIMENS 79 aeruginosus Morelet, Unio : 1849, Testacea Novissima, 1 : 29 (rivulo Michol, circa Palenquea- num vicum [Chiapas, Mexico]). Measured holotype BMNH 93.2.4.2019, figured by Fischer and Crosse, 1894, Miss. Sci. au Mexique, pt. 7, 2 : 596, pi. 62, figs. 2, 2a, 2b ; paratype BMNH 93.2.4.2020. aethiopiformis "Ihering" Simpson, Unio: 1914, Descr. Cat. Naiades, 3: 1312 [nomen nudum]. Specimens under this name, BMNH 1891.4.13.51-55. Distributed by Ihering but never described by him. aferula Lea, Unio : 1864, Proc. Acad. Nat. Sci. Phila., 16 : 109 (Lake Nyassa, Central Africa, John Kirk) ; 1866 Jour. Acad. Nat. Sci. Phila. (2) 6 : 34, pi. 13, fig. 34 ; 1867, Obs. Unio, 11 : 38. Holotype USNM 84058 ; specimen from Kirk's lot, but not seen by Lea, BMNH 64.5.14.2. alata Sowerby, Hyria : 1869, Conch. Iconica, 17, Hyria, pi. 5, species 13 (Guayana). Holotype BMNH 1849.6.1.2 ; 2 paratypes BMNH 1849.6.1.2 1-2. alata Lea, Spatha : 1864, Proc. Acad. Nat. Sci. Phila., 16 : 109 (Lake Nyassa, Central Africa, John Kirk) ; 1866, Jour. Acad. Nat. Sci. Phila. (2) 6 : 35, pi. 12, fig. 31 ; 1867, Obs. Unio, 11 : 39. Holotype USNM 86776 ; 2 specimens from Kirk's lot, but not seen by Lea, BMNH 62.9.25.4. ambiguus " Parreyss " Philippi, Unio : 1847, Abb. und Besch. Conch., 3 : 7 [47], pi. 3, fig. 2. (Nova Holandia, purchased from Parreyss). Holotype BMNH 41.4.29.103 teste and figured by McMichael and Hiscock, Australian Jour. Marine and F. W. Res., 9, pi. i, figs. 1-2. While there is no evidence that this specimen was the one figured by Philippi, it is probably as close to an authentic type as can be found. angasii Sowerby, Anodon : 1870, Conch. Iconica, 17, Anodon, pi. 32, species 127 (South Australia [Strangway's River, North Australia]). [Credited to Lea Obs., 12 but not described by him], Holotype BMNH 70.10.26.41 refigured by McMichael and Hiscock, 1958, Australian Jour. Marine and F. W. Res., 9 : pi. 3, figs. 5-6. angustior Hanley and Theobald, Unio generosus : 1872, Conch. Indica, p. 22, pi. 46, fig. 7 (Pegu) Holotype BMNH 1968655 annulatus Sowerby, Anodon : 1867, Conch. Iconica, 17, Anodon, pi. 17, species 67 (Hab.?, Cuming colln.). BMNH [lost.] aplatus Reeve, Unio : 1865, Conch. Iconica, 16, Unio, pi. 28, species 143 (Island of Chiloe, Chili, Cuming colln.). BMNH [lost]. arcuans Fischer and Crosse, Unio calamitarutn : 1894, Miss. Sci. au Mexique, pt. 7, 2 : 613, pi. 64, figs. 5, 5a [Mexico, Morelet colln]. Figured holotype BMNH 93.2.4.2009. areolatus Sowerby, Anodon : 1867, Conch. Iconica, 17, Anodon, pi. 10, species 28 (Hab.?, Cuming colln.). BMNH [lost]. askewi Marsh, Unio : 1896, Nautilus, 10 : 92 (Village Creek, Hardin Co., Texas ; Sabine River, Texas) ; Marsh 1897, Nautilus, 10 : n, pi. i, figs. 3-4. Figured holotype ANSP 7O448a ; 2 paratypes BMNH 98.2.1.36-37, from the Sabine River, Texas. auklandicus Gray, Unio : 1843 [in] Dieffenbach, Travels in New Zealand, 2 : 257 (Bay of Islands ; Auckland in the Bay of Amabrusa) restricted by McMichael and Hiscock 1958, Australian Jour. Marine and F. W. Res., 9 : 455 to near Auckland, North Island, New Zealand. Holotype BMNH 1951.9.6.17 figured Ibid., pi. 12, figs. 13-14, is two unmatched valves original no. 1842.11.5.75. auratus Reeve, Unio : 1865, Conch. Iconica, 16, Unio, pi. 29, species 150 (Island of Chiloe, Chili, Cuming colln.). Holotype BMNH 1965144 ; 2 paratypes BMNH 1965145. auratus Sowerby, Unio : 1866, Conch. Iconica, 16, Unio, pi. 45, species 245 (South America, Hanley colln.). [not in BMNH]. avae Theobald, Monocondylaea : 1873, Jour. Asiatic Soc. Bengal, 42 (2) : 209, pi. 17, fig. 5 (Mandelay regno Birmanica). Figured holotype BMNH 88.12.4.1743. baikii A. Adams, Spatha : 1866, Proc. Zool. Soc. London, p. 447 (River Niger [Nigeria]). Of the three syntypes of this species in the BMNH, one from the Cuming colln., is closest to Adams' original measurements and is, here selected, lectotype BMNH 196466, plate i fig. 3. Length 116, height 75, width 46 mm ; paralectotype BMNH 196467, Cuming colln. ; para- lectotype BMNH 78.1.28.199. ex. A. Adams. 8o R. I. JOHNSON bakeri, H. Adams, Unio : 1866, Proc. Zool. Soc. London, p. 376 ([Lake] Albert N'yanza [Central Africa]). Holotype BMNH 1867.1.9.1, figured by Smith, 1892. Ann. and Mag. Nat. Hist. (6) 10, pi. 12, fig. ii, consists of one valve. bambousearum Morelet, Anodon : 1851, Testacea Novissima, 2 : 24 (rivulos prope vicum Palenqueanum, prov. Chiapas [Mexico]). Measured holotype BMNH 1965146, figured by Fischer and Crosse, 1894, Miss. Sci. au Mexique, pt. 7, 2 : 527, pi. 63, figs. 6, 6a. bellua Morelet, Anodonta : 1866, Rev. etMag. deZool. (2) 18 : 167 (lacu Toui-Sap, [Cambodia]). Lectotype MCZ 175610 selected by Johnson, 1956, Bull. Mus. Comp. Zool. 115 : 107, pi. i, fig. i ; paralectotype BMNH 1965147. bengalensis Sowerby, Anodon : 1867, Conch. Iconica, 17, Anodon, pi. 14, species 49 (Bengal Rivers, Cuming colln.). BMNH [lost]. bhamoensis Theobald, Unio : 1873, Jour. Asiatic Soc. Bengal, 42 (2) : 207, pi. 17, fig. i (prope Bhamo, Regno Birmanico ; necnon in Prome occidentali Provincia Pegu). Figured holotype BMNH 88.12.4.1672, refigured by Hanley and Theobald, 1876, Conch. Indica, p. 62, pi. 155, fig. 2 ; paratype BMNH 88.12.4.1673. bicaelatus Reeve, Unio : 1865, Conch. Iconica, 16, Unio, pi. 26, species 130 (Hab. ?, Cuming colln.). BMNH [lost]. bischoffi " Ihering " Simpson, Unio: 1914, Descr. Cat. Naiades, 3: 1312 [nomen nudum]. Specimens under this name, BMNH 1891.4.13.22—23, distributed by Ihering but never dis- cribed by him. bonelli Sowerby, Anodon : 1867, Conch. Iconica, 17, Anodon, pi. n, species 34 (Lago Mag- giore [Italy] Cuming colln.). Holotype BMNH 1964379. breviculus Call, Unio : 1887, Proc. United States Natl. Mus., 10 : 499, pi. 28 (Currant River, Shannon Co., Missouri and in Jack's Fork and Big Creek, tributaries of it). Holotype MCZ 5020 ; 2 paratypes BMNH 98.2.1.3940 from Big Creek. brevis Sowerby, Anodon : 1870, Conch. Iconica, 17, Anodon, pi. 31, species 124 (Rio Plata). Mentioned as in BMNH [lost]. burtoni Woodward, Unio : 1859, Proc. Zool. Soc. London, 27 : 349, pi. 47, fig. i (Lake Tangan- yika [Central Africa]). Holotype BMNH 1859.12.23.9, refigured by Sowerby 1866, Conch. Iconica, 16, Unio, pi. 47, species 251 ; paratype BMNH 1965220. calamitarum Morelet, Unio : 1849, Testacea Novissima, 1 : 30 (rivulum Baluntie, propre Palenqueanum vicum [Chiapas, Mexico]). Not figured by Fischer and Crosse, 1894, Miss. Sci. au Mexique, pt. 7, 2. Morelet gives the measurements of the type as : Altit. 29, latit. 51, diam. 16 mm. [not found]. Lectotype, here selected, BMNH 1893.2.4.2010, plate 2 fig. 5 Length 54, height 32, width 26 mm. callifera Martens, Anodonta : 1860, Proc. Zool. Soc. London, 28 : 15 (Siam). Holotype BMNH 59.8.1.20, figured by Haas 1913, [in] Martini and Chemnitz, Conch. Cab. (2) 9, pt. 2, sec. 2, pi. 46, fig- 3- cambojensis Sowerby, Unio : 1866, Conch. Iconica, 16, Unio, pi. 42, species 231 (Camboja, Cuming colln.). Holotype BMNH 1965148 consists of one valve ; 2 paratypes BMNH 1965149. carinthiacus Ziegler, Unio : 1835 [in] Rossmassler, Icon. Land- und Susswasser Moll, (i) 1 : 21 ; Ibid., 3 : 30, pi. 15, fig. 209 (Carinthia). Paratype BMNH 1965216, figured by Reeve, 1856, Conch. Iconica, 16, Unio, pi. 30, species 157 [credited to Ziegler in index]. carolinensis Sowerby, Castalia : 1869, Conch. Iconica, 17, Castalia, pi. 2, species 6 (South, Carolina). Holotype BMNH 1841.4.6.149. championi Martens, Unio : 1900, Biol. Cent. Americana, Moll., p. 508, pi. 38, fig. 9, ga (W. Guatemala : Paso Antonio, in the Pacific coast-region). Paratypes BMNH 1901.6.22. 1538-9. charruana Orbigny, Unio : 1835, Mag. de Zool., p. 35 (Banda orientali, republica Uruguayensi orientali) ; 1846, Voy. Amer. Merid., 5, pt. 3, p. 606, pi. 71, figs. 8-14 (tous les ruisseaux et les petites rivieres dupuis Maldonado, Montevideo, jusqu'a Las Bacas). Holotype BMNH 1854.9. 4.15 from Rio Rosario ; 12 paratypes BMNH 1854.9.4.15/1-12. cheeziana Sowerby, Anodon : 1867, Conch. Iconica, 17, Anodon, pi. 15, species 52 (Hab. ?, Cuming Colin.). Holotype BMNH 1964399. UNIONACEA: TYPES AND FIGURED SPECIMENS 81 chilensis " Parreysis " Sowerby, Unio : 1867, Conch. Iconica, 16, Unio, pi. 56, fig. 286 (Chili, Taylor colln.). Figured holotype BMNH 74.12.11.11. chinnerethensis Preston, Unio. 1913, Jour, and Proc. Asiatic Soc. Bengal (n.s.) 9 : 473, pi. 27, fig. 10, loa (Lake of Tiberias, Galilee). Paratypes BMNH 1913.7.30.18 ; 1914.1.7.203-205. chiquitana Orbigny, Anodonta : 1835, Mag. de Zool., p. 41 (Corrientes, republica Argentina ; provincia Chiquitensi, rep. Boliviana). New name for Anodon trigonum Spix, pi. 22, fig. 2 ; 1846, Voy. Amer. Merid., 5, pt. 3, listed as a synonym of Anodonta trigona Spix. ciconia Gould, Anodon : 1851, Proc. Boston Soc. Nat. Hist., 4 : 92 (Mexico?). Holotype MCZ 189084, figured by Johnson, 1964, Bull. United States Natl. Mus., 239, p. 54, pi. 34, fig. i ; paratype BMNH 1964397 figured by Sowerby, 1870, Conch. Iconica, 17, Anodon, pi. 29, species ii5a. Species H5b BMNH 1964389 is not this species. clappertoni Koenig, Anodon : 1826 [in] Denham and Clapperton, " Narrative of Trav. and Discov. in N. and Centr. Africa, Appendix ", p. 255 (Gammaroo River [appears to be a locality called Gambaroo on the Komadugu Yobe River, also called the Yaou or Yo]). Syntype BMNH [not catalogued]. cochlearis Sowerby, Anodon : 1870, Conch. Iconica, 17, Anodon, pi. 33, species 135 (Hab. ?). BMNH [lost]. cocoduensis "White" Reeve, Unio : 1865, Conch. Iconica, 16, Unio, pi. 24, species 117 (Hab.?, Cuming colln.). Holotype BMNH 1965187 ; paratype BMNH 1965188. compressa Martens, Spatha : 1860, Proc. Zool. Soc. London, 28 : 16 (Khaokho, NE of Pak- priau, Siam). Holotype BMNH 59.8.1.21, figured by Haas 1913, [in] Martini and Chemnitz, Conch. Cab. (2)9, pt. 2, sec. 2, pi. 36, fig. 2. contorta Lea, Triquetra : see under, lanceolata Lea, Triquetra. cor data Sowerby, Cast alia : 1869, Conch. Iconica, 17, Castalia, pi. 2, species 8 (British Guayana, Cuming colln.). Holotype BMNH 1965197 ; 2 smaller paratypes BMNH 1965198. coreanus Martens, Unio : 1886, Sitzber. Gesell. Natur. Freunde, Berlin, p. 70 (Fluss Hangang, 15 km. oberhalb Soul [Prov. Kyongkwido] Korea) ; Martens, 1905, Zool. Jahrb., Suppl. 8, p. 50, pi. 3, fig. 5. Paratype BMNH 94.11.23.10. corium Reeve, Unio : 1864, Conch. Iconica, 16, Unio pi. 10, species 39 (Chiapa, Mexico, Cuming colln.). Holotype BMNH 1965209. corrientesensis Orbigny, Monocondylaea : 1835, Mag. de Zool., p. 38 (Rio Corrientes, pro- vincia Corrientesensi, republica Argentina) ; 1846, Voy, Amer. Merid., 5, pt. 3, p. 613, pi. 68, figs. 8-10 (le Rio Batel, au sud de la province de Corrientes, Argentine). Holotype BMNH 1854.9.4.28 ; 3 paratypes BMNH 1854.9.4.28/1-3. crepera Lea, Anodonta : 1851 Proc. Zool. Soc. London for 1850, pt. 18, p. 198 (Bongabon, Luzon, Philippine Islands, Cuming) ; Lea, 1851, Ann. and Mag. Nat. Hist., 8 : 494 ; Lea, 1860, Jour. Acad. Nat. Sci. Phila. (2) 4 : 238, pi. 23, fig. 117 ; Lea 1860, Obs. Unio, 7 : 56. Lea saw five or six specimens. Holotype BMNH [lost] ; 2 paratypes BMNH 1965158 ; paratypes USNM 86602. The two specimens in the BMNH 42.5.10.1538 labeled Saul, Luzon, are not types. crispata Gould, Unio : 1843, Proc. Boston Soc. Nat. Hist., 1 : 141 ([Tavoy] British Burmah [Burma]). Lectotype MCZ 186099, selected by Johnson, 1964, Bull. United States Natl. Mus., 239, p. 62, pi. 32, fig. 3 ; paralectotype BMNH 1965223. crispisulcatus Benson, Unio : 1862, Ann. and Mag. Nat. Hist., 10 : 193 ; Sowerby, 1866, Conch. Iconica, 16, Unio, pi. 49, fig. 262 (Hab.?, Cuming colln.). BMNH 1964358. Though credited to Lea ms., this specimen is probably a type. crocodilorum Morelet, Unio : 1849, Testacea Novissima, 1 : 28 (flumen Usumasinta [Guate- mala]). Measured holotype BMNH 93.2.4.2030, figured by Fischer and Crosse 1894, Miss. Sci. au Mexique, pt. 7, 2 : 577, pi. 60, fig. 3 ; 2 paratypes BMNH 93.2.4.2031-32, were also figured, pi. 60, figs. 4—5 and were named varieties semipustulata and praestricta respectively. cryptoradiata Putzeys, Spatha : 1898, Ann. Soc. Mai. Belgique, 33 ; Bull. Seances, p. xxiv, figs. 14-15 (le Stanley- Pool, pres de Leopold ville [Congo]). 2 paratypes BMNH 1902. 1 1. 20. i— 2, from Putzeys, purchased from H. B. Preston. 82 R. I. JOHNSON cumingii Lea, Anodonta : 1850, Proc. Zool. Soc. London, 18 : 199 (Malacca, Cuming colln.) ; cumingii Lea, Monocondylaea, 1860, Jour. Acad. Nat. Sci. Phila. (2) 4 : 235, pi. 33, fig. 114 ; 1860, Obs. Unio, 7 : 53. Holotype USNM 86350 ; paratype BMNH [not catalogued] figured by Sowerby, 1870, Conch. Iconica, 17, Anodon, pi. 31, species 122. cumingii Dunker, Galatea ; 1860, [in] Bernardi, Monog. Galatea et Fischeria, p. 35, pi. 6, figs. 7-8 ; pi. 9, fig. 8 (le Gabon, Cuming colln.). BMNH [lost]. cumingii Lea, Unio : 1852, Proc. Acad. Nat. Sci. Phila., 6 : 54 (Northern part of China, H. Cuming) ; Lea 1860, Jour. Acad. Nat. Sci. Phila. (2) 4 : 240, pi. 35, fig. 120 ; Lea 1860, Obs. Unio, 7 : 58. Holotype USNM 83897 ; paratype BMNH 1964360 figured by Sowerby, 1866, Conch. Iconica, 16, Unio, pi. 49, species 264. cuneata Preston, Mutela : 1910, Ann. and Mag. Nat. Hist. (8) 6 : 62, pi. 5, fig. 13 (Karonga, northern end of Lake Nyassa [Central Africa]). Paratype BMNH 1910.9.28.9. dactylus Sowerby, Anodon : 1867, Conch. Iconica, 17, Anodon, pi. 19, species 75 (Hab. ?, Cuming colln.). BMNH [lost]. dactylus Morelet, Unio : 1845, Moll, de Portugal, p. no, pi. 14, fig. 2 (affluent de la Guadiana pres de Castro- Verde en Algarve). Holotype BMNH 93.2.4.1587. dahomeyensis Lea, Anodonta : 1859, Proc. Acad. Nat. Sci. Phila., 11 : 154 (Dahomey, West Africa, Mr. Frazer, Cuming colln.) ; Lea 1859, Jour. Acad. Nat. Sci. Phila. (2) 4 : 261, pi. 41, fig. 141 ; Lea 1860, Obs. Unio, 7 : 79 ; Sowerby, 1870, Conch. Iconica, 17, Anodon, pi. 36, species 151. Holotype BMNH 1965163. The type lot also contained two smaller speci- mens BMNH 1965164 not seen by Lea. dalyi E. A. Smith, Mulleria : 1898, Proc. Mai. Soc. London, 3 : 14, text figs. (Probably near Mudgiri, Kadur District, Mysore Dist., Southern India). Lectotype BMNH 1897.11.19.1, selected by Pain and Woodward 1961, Jour, of Conch., 25 : 5, specimen in upper two drawings ; 2 paralectotypes BMNH 1897.11.30.19—20. delessertii Bernardi, Fischeria : 1860, Monog. Galathea et Fischeria, p. 46, pi. 3, figs. 3-4 ; pi. 9, fig. 5 (Les cours d'eau pres du cap Palmas, GumSe, Bernardi [Paris Mus.?] et Cuming colln.). BMNH [lost]. delphinulus Morelet, Unio : 1849, Testacea Novissima, 1 : 31 (paludosa flum. Usumasinta et lacum Petenensem [lac de Flores ou Tha, Guatemala]). The specimen figured by Fischer and Crosse, 1894, Miss. Sci. au Mexique, pt. 7, 2 : 557, pi. 63, figs. 2, 2a, 2b in the Paris Museum may be a type, but it is larger than the measured holotype. Probable measured holotype BMNH 93.2.4.1588. Morelet's description gives Alt. 26, Lat. 60, Diam. 14 mm. The prob- able holotype measures, without the " wing ", Length 29, height 60, width 14 mm. The label has the additional data given above in brackets. dembeae Reeve, Unio : 1865, Conch. Iconica, 16, Unio, pi. 29, species 153 (Dembea, Abyssinia, Cuming colln.). BMNH [lost]. demeraraensis Lea, Unio : 1859, Proc. Acad. Nat. Sci. Phila., 11 : 152 (Demerara, British Guiana, Cuming colln.) ; Lea, 1860, Jour. Acad. Nat. Sci. Phila. (2) 4 : 253, pi. 39, fig. 133 ; Lea, 1860, Obs. Unio, 7 : 71. Lea only saw the holotype, which was not found in the BMNH, or in the USNM. dernaica Pallary, Margaritana : 1928, Jour, de Conch., 72 : 22, pi. 4, figs. 1-2 (1'oued Derna, affluent de la rive gauche de 1'oum er Riba, a la hauteur de Tagnzirt (moyen Atlas, meridional) region de Tadla). Paratype BMNH 1937.12.30.13022. deviatus "Anthony" Reeve, Unio : 1964, Conch. Iconica, 16, Unio, pi. 15, species 61 (North America, Cuming colln.). Holotype BMNH 1965210, with Anthony's original label to H. Cuming, " Tennessee, very rare, best I have. " digitatus Morelet, Unio : 1851, Testacea Novissima, 2 : 24 (flumen Usumasinta [Peten Prov., Guatemala]). Measured holotype BMNH 93.2.4.2035, figured by Fischer and Crosse, 1894, Miss. Sci. au Mexique, pt. 7, 2 : 563, pi. 60, fig. i. digitiformis Sowerby, Unio : 1868, Conch. Iconica, 16, Unio, pi. 65, species 333 (India, Taylor colln.). Holotype BMNH 1965199. • diminutus Lea, Unio : 1 859, Proc. Acad. Nat. Sci. Phila., 11:151 (East Africa, H. Cuming and S. Hanley) ; Lea, 1806, Jour. Acad. Nat. Sci. Phila. (2) 4 : 254, pi. 39, fig. 134 ; Lea, 1860, Obs. UNIONACEA: TYPES AND FIGURED SPECIMENS 83 Unio, 7 : 72. Holotype BMNH 1965165 Cuming Colin. The type lot also contains a smaller specimen BMNH 1965166, not seen by Lea. dolabe.Ua Sowerby, Castalia : 1869, Conch. Iconica, 17, Castalia, pi. 3, species 13, figs, a-c (Hab. ?, Cuming colln.). Holotype BMNH 1965185 [River Amazon] ; paratype BMNH 1965186. dugasti Morlet [sic], Unio : 1892, Jour, de Conch., 40 : 86 (riviere Outhene, petit affluent du M6kong, Laos) ; 1893, Ibid., 41 : 156, pi. 6, fig. 4, 2 paratypes BMNH 93.12.8.137-8, ex. Ph. Dautzenberg. durrovensis Phillips, Margaritifera : 1928, Proc. Mai. Soc. London, 18 : 72, pi. 3, fig. i ; pi. 4, fig. i (River Nore, Durrow, Queen's Co., England). 2 figured syntypes BMNH 1928.10.23. 1-2. dysonii Lea, Unio : 1859, Proc. Acad. Nat. Sci. Phila., 11 : 152 (Honduras, D. Dyson, Cuming colln.) ; Lea, 1860, Jour. Acad. Nat. Sci. Phila. (2) 4 : 252, pi. 39, fig. 132 ; Lea, 1860, Obs. Unio, 7 : 70. Holotype BMNH 1965173. The type lot also contains a smaller specimen BMNH 1965174, not seen by Lea. electrinus Reeve, Unio : 1865, Conch. Iconica, 16, Unio, pi. 25, species 121 (Hab. ?, Cuming colln.). BMNH [lost]. elliottii Lea, Unio : 1856, Proc. Acad. Nat. Sci. Phila., 8 : 262 (Othcalooga Creek, Gordon Co., Georgia, Elliott) ; Lea, 1858, Jour. Acad. Nat. Sci. Phila. (2) 4 : 54, pi. 5, fig. 37 ; Lea, 1858, Obs. Unio, 6 : 54. Holotype USNM 84019 ; paratype BMNH 1965160, Cuming colln. ex Lea, figured by Reeve, 1864, Conch. Iconica, 16, Unio, pi. 3, species 20. elongata Longstaff, Nodularia (Caelatura) parreysii : 1914, Jour. Linn. Soc. London, 32 : 255, pi. 18, figs. 9-10 (White Nile River at Ad-Duwem, Tawlla, and Gebel Ahmad Agha, Southern Sudan). Figured holotype BMNH 1923.6.8.1218 from Gebel Admad Agha. episcopalis Tristram, Unio : 1865, Proc. Zool. Soc. London, p. 544 (Orontes River, Palestine). Lectotype, here selected, BMNH 1936.3.10.3. plate 2 fig. i . Length 80, height 47, width 28 mm. esula Orbigny, Iridina : 1835, Mag. de Zool. p. 43, nomen nudum ; 1843, Voy. Amer. Merid., 5, pt. 3, p. 597 (tous les lacs du centre de la Bolivia ; pays des Guarayos, province de Chiquitos, et dans toute la province de Moxos). The type was not figured, and since it was not found in the BMNH, it is presumed that it was lost between the time of description and the preparation of the plates. eurhynchus "Bronn" Kuester, Unio : 1861 [hi] Martini and Chemnitz, Conch. Cab. (2) 9, pt. 2, p. 237, pi. 79, fig. 5 ([Minas Geraes] Brasilien). BMNH 1965217, distributed by Bronn before description by Kuester. evansi Adams and Angus, Unio : 1864, Proc. Zool. Soc. London, p. 39 (Lagoons of Lower Murray R., South Australia). Holotype BMNH 1870.10.26.42, figured by McMichael and Hiscock, 1958, Australian Jour. Marine and F. W. Res., 9, pi. i, figs. 6-7. exasperata Sowerby, Hyria : 1869, Conch. Iconica, 17, Hyria, pi. 2, species 3 (British Guayana). BMNH [lost]. exilis Morelet, Monocondylus : 1866, Jour, de Conch., 14 : 63 (in torrentibus montanis Cam- bodiae [lac Touli-sap au Combodje]). Holotype BMNH 93.2.4.1981, figured by Morelet, 1875, Series Conch., pt. 4, p. 340, pi. 17, fig. i. exoticus Sowerby, Anodon : 1867, Conch. Iconica, 17, Anodon, pi. 16, species 57 (Rio de la Plata [Argentina]). Holotype BMNH 1965207, consists of one valve. explicatus Morelet, Unio : 1849, Testacea Novissima, 1 : 28 (flumen Usumasinta, ad pagum Balancan Tabascensium [Mexico]). Measured holotype BMNH 93.2.4.2027, figured by Fischer and Crosse, 1894, Miss. Sci. au Mexique, pt. 7, 2 : 594, pi. 61, fig. i. expressa Martens, Anodonta : 1900, Natur. Deutschen Mai. Ges., 32 : 12 (Lake Danau-Baru, Indragiri, Sumatra). Two syntypes are figured by Haas 1920, [in] Martini and Chemnitz, Conch. Cabinet 9, pt. 2, no. 2, p. 304, pi. 38, figs. 1-2 ; syntype BMNH 1901.6.14.3. exulceratus "Porro" Sowerby, Anodon : 1870, Conch. Iconica, Anodon, 17, pi. 33, species 131. Figured holotype BMNH 1841.5.6.127, from Zeigler. faba Orbigny, Unio : 1835, Mag. de Zool., p. 35 (Banda orientali, republica Uruguayensi 84 R. I. JOHNSON orientali). Is a variety of Unio charruana Orb., teste, Orbigny, 1846, Voy. Amer. Merid., 5, pt. 3, p. 606. 5 syntypes BMNH 1854.12.4.839. falcatus Higgins, Mycetopus : 1868, Proc. Zool. Soc. London, p. 179, pi. 14, fig. 6 (Forest streams near Chyavetas, Upper Amazons [Brazil], E. Bartlett). Holotype BMNH 68.4.3.6 ; paratype MCZ 74218, ex. R. F. Geale. favidens Benson, Unio : 1862, Ann. and Mag. Nat. Hist. (3) 10 : 188 (Ganges [River], between Cawnpore and Allahabad [India]). 2 syntypes BMNH 1965215 from Benson ex. Cuming colln. The type figured by Reeve, 1865, Conch. Iconica, 16, Unio, pi. 26, species 131 Cuming colln. BMNH [lost]. ferrarisii Orbigny, Anodonta : 1835, Mag. de Zool., p. 40 (Banda orientali [Rio del Rosario], Uruguayensi orientali) ; 1846, Voy. Ame'r. Merid., 5, p. 615, pi. 74, fig. 3. Listed as synonym oiAnodontes sirionos Orbigny. Figured holotype BMNH 1854.9.4.34, consists of one valve. flgdiana Bloomer, Caelatura aegyptiaca : 1946, Proc. Mai. Soc. London, 27 : 69, pi. 6, fig. 3 (Wad Figda, canal between Khartoum and Sennaar, Soudan). Paratype BMNH 1948.5.58. fimbriata Frierson, Larnpsilis : 1907, Nautilus, 21 : 86, pi. 12, two upper figs, and lower left hand fig. (Valles River [San Luis Potosi Prov.] Mexico). Figured holotype Mus. Zool., Univ. Michigan 191161 ; 2 paratypes BMNH 1910.9.30.170-171. fluctiger, Lea, Unio : 1859, Proc. Acad. Nat. Sci. Phila., 11 : 152 (Hab. ?, H. Cuming) ; Lea, 1860, Jour. Acad. Nat. Sci. Phila. (2) 4 : 250, pi. 39, fig. 130 ; Lea, 1860, Obs. Unio, 7 : 68. Refigured by Sowerby, 1866, Conch. Iconica, 16, Unio, pi. 42, species 299. Holotype BMNH 1965169. The type lot also contained a smaller specimen, BMNH 1965170 not seen by Lea. fontaineana Orbigny, Unio : 1835, Mag. de Zool., p. 36 (Rio Parahiva, imperio Brasiliano) ; 1846, Voy. Amer, Mdrid., 5, pt. 3, p. 605, pi. 69, figs. 6-7. Holotype BMNH 1854.9.4.49 ; paratype BMNH 1854.9.4.49/1. footei Theobald, Unio : 1876, Jour. Asiatic Soc. Bengal, 45 (2) : 187, pi. 14, fig. 9 (Kistna flumine prope " Gutparba falls "). Figured holotype BMNH 88.12.4.1651. fossiculifera Orbigny, Monocondylaea : 1835, Mag. de Zool., p. 38 (Rio Parana, provincia Corrientesensi, republica Argentina) ; 1846, Voy. Ame'r. Merid., 5, pt. 3, p. 614, pi. 80, figs. 5-7 (Parana, a Iribucua, distant de vingt lieues au-dessus de corrientes, republique Argentine). Holotype BMNH 1854.9.4.1 ; 7 smaller paratypes BMNH 1854.9.4.1/1-8. fragilis Sowerby, Unio : 1865, Conch. Iconica, 16, Unio, pi. 30, species 155 (Island of Chiloe, Chili, Cuming colln.). Non Unio fragilis Swainson, 1823, changed to Diplodon cuprinus Simpson 1900, Proc. United States Natl. Mus., 22 : 883. Figured holotype BMNH [not catalogued] ; 2 paratypes [not catalogued]. fragilis Hanley and Theobald, Unio corrugatus : 1872, Conch. Indica, p. 21, pi. 45, fig. 4 (no locality). Figured holotype BMNH 196851. framesi Connolly, Indonaia : 1925, Records Albany [South Africa] Mus., 3 : 265, pi. 12, figs, i, 4 (Transvaal : near Premier Mine, Pretoria Dist., South Africa). Holotype Senckenberg Mus.; paratypes BMNH 1926.7.7.8 and 1937.12.30.128. frier soni Wright, Unio : 1896, Nautilus, 9 : 134, pi. 3 (Bayou Pierre, an arm of the Red River in DeSoto Parish, Louisiana). Lectotype USNM 133432, selected by Johnson, 1967, Occ. Papers on Moll., 3 : 6, pi. 5, fig. 4 ; 4 paralectotypes BMNH 98.2.1.43-44 and 05.8.15.38-39. fuligo Reeve, Unio : 1865, Conch. Iconica, 16, Unio, pi. 30, species 159 (Hab. ?, Cuming colln.). BMNH [lost]. fulmineus " Parreyss " Philippi, Unio: 1847, Abb. und Besch. Conch., 3 : 6[46], pi. 3, figs. 5-6 (Nova Hollandia). 3 probable syntypes BMNH 41.4.18.135-138, from Parreyss. gibba Sowerby, Anodon : 1867, Conch. Iconica, 17, Anodon, pi. 6, species 13 (River Kiang, [Vietnam] Cuming colln.). Holotype BMNH 1964372. gracilis Lea, Anodonta : 1851, Proc. Zool. Soc. London for 1850, pt. 18, p. 197 (Dingle, Island of Panay, Philippine Islands, Cuming colln.) ; Lea, 1851, Ann. and Mag. Nat. Hist., 8 : 193 ; Lea, 1860, Jour. Acad. Nat. Sci. Phila. (2) 4 : 239, pi. 34, fig. 119 ; Lea, 1861, Obs. Unio, 7 : 57. Holotype USNM 86722 ; paratype BMNH [not found], figured by Sowerby, 1857, Conch. Iconica, 17, Anodon, pi. 14, species 45 ; 3 paratypes BMNH 1968657. UNIONACEA: TYPES AND FIGURED SPECIMENS 85 gracilis "Parreyss" Martens, Iridina : 1866, Mai. Blatt., 13 : n [nomen nudum, listed as a synonym of Spathia rostrata Rang] BMNH 41.4.28.152, distributed by Parreyss before description by Martens. graueri Haas, Caelatura : 1927, Senckenbergiana, 9 : 21 (Urwald Ukaika, norwestlich vom Albert— Edward— See, Innerafrika) . Paratype BMNH 1937.12.30.13117. grijalvae Morelet, Andonta : 1884, Jour, de Conch., 32 : 123 (Rio dos Idolos, un des bras du fleuve Grijalva, Tabasco [Mexico]). Lectotype, here selected, BMNH 93.2.4.2009, figured by Fischer and Crosse, 1894, Miss. Sci. au Mexique, pt. 7, 2 : 532, pi. 59, fig. i, consists of the opposite valve of the specimen figured by Fischer and Crosse and is somewhat smaller than the original measured type. guaraniana Orbigny, Unto : 1835, Mag. de Zool., p. 37 (Rio Parana, provincia Corrientesensi, republica Argentina) ; 1846, Voy. Amer. Merid., 5, pt. 3, p. 608, pi. 69, figs. 10-12 (pres du village d'ltaty, Corrientes, lorsque le Parana, [Argentina]). Measured holotype BMNH 1854.12.4.841, the figure is enlarged ; paratype BMNH 1854.12.4.841/1, consists of one valve. guarayana Orbigny, Monocondylaea : 1835, Mag. de Zool., p. 38 (Rio San Miguel, Guarayos [Chiquitos], Boliviana) ; 1846, Voy. Amer. Merid., 5, pt. 3, p. 614, pi. 68, figs. 4-7. Holotype BMNH 1854.9.4.17 ; paratypes BMNH 1854.9.4.17/1-3. guatybae "Ihering" Simpson, Unio : 1914, Descr. Cat. Naiades, 3 : 1312 [nomen nudum]. Specimens under this name, BMNH 1891.14.13.34-37 were distributed by Ihering but never described. gubernaculum Reeve, Unio : 1865, Conch. Iconica, 16, Unio, pi. 28, species 146 (Hab. ?, Cuming colln). Holotype BMNH 1965203 ; paratype BMNH 1965204. guppyi E. A. Smith, Unio : 1885, Proc. Zool. Soc. London, p. 608, pi. 37, fig. 88a, b (Shortland Island, Solomon Islands). Figured holotype BMNH 1885.11.3.461 ; paratypes BMNH 1885.11.3.462-5. hanleyana Sowerby, Cast alia : 1869, Conch. Iconica, 17, Castalia, pi. i, species 5 (Hab. ?, Hanley colln.). Holotype BMNH 1900.3.19.5. hargeri E. A. Smith, Mutela : 1908, Proc. Mai. Soc. London, 8 : 14, text fig. (Lake Mweru, Central Africa). Holotype BMNH 1907.11.11.55, consists of the one valve on which the description was based. harlandi Baird and H. Adams, Unio : 1867, Proc. Zool. Soc. London, p. 492, pi. 26, fig. 3, 3a (Shanghai, North China). Figured holotype BMNH 1965222. herculeus Sowerby, Anodon : 1867, Conch. Iconica, 17, Anodon, pi. 3, species 7 ([Tago, Japan] Cuming colln.). Holotype BMNH 1965200, consists of one valve. hermosus Bourguignat, Unio : 1889, Bull. Soc. Mai. de France, 6 : 38, new name for Unio nyassaensis, var. E. A. Smith 1881, Proc. Zool. Soc. London, p. 298, pi. 34, fig. 34b (Lake Nyassa). Holotype BMNH 1880.12.20.130, consists of one valve ; 2 paratypes BMNH 1880.12.20.131. Mans Sowerby, Anodon : 1867, Conch. Iconica, 17, Anodon, pi. 4, species 8 (Hab. ?, Cuming Colin.). Holotype BMNH 1965205 ; paratype BMNH 1965206. horda Gould, Anodon : 1855, Proc. Boston Soc. Nat. Hist., 5 : 299 (Comanche Creek, [Tribu- tary of the Llando River, close to the present site of Mason, Mason Co.], Texas [teste Taylor, 1967, Veliger, 10 : 153]). Measured holotype USNM 678301 ; paratype BMNH 196465, figured by Sowerby, 1867, Conch. Iconica, 17, Anodon, pi. 18, species 66, Cuming colln. horei E. A. Smith, Unio : 1880, Ann. and Mag. Nat. Hist. (5) 6 : 429 (Lake Tanganyika [Central Africa]) ; 1881, Proc. Zool. Soc. London, p. 299, pi. 34, fig. 37. Holotype BMNH 80.12.20.42. humilior Martens, Unio scutum : 1899, Archiv. fur Natur., 1 : 45, pi. 5, fig. i (Chindwinfluss bei Kalewa und in einem Nebenfluss desselben, dem Yufluss [Burma]). 4 paratypes BMNH 99.6.21.6-9. hylaea Orbigny, Unio : 1835, Mag. de Zool., p. 36 (provincia Chiquitensi, republica Boliviana) ; 1846, Voy. Amer. Merid., 5, pt. 3, p. 607, pi. 69, figs. 8-9 (Santa Cruz de la Sierra et de Chi- quitos en Bolivia, principalement dans les rivieres nominees Palometas, Pari, et Tucabaca). Holotype BMNH 1854.12.4.843, from Chiquitos ; paratype BMNH 1854.12.4.843/1, consists 86 R. I. JOHNSON of one valve. Paratypes BMNH 1854.12.4.842 consists of two complete specimens and three odd valves from Santa-Cruz. impressa Anthony, Alasmodon : 1865, Amer. Jour. Conch., 1 : 157, pi. 12, fig. 4 (Tennessee). Holotype MCZ 150666 ; 3 paratypes BMNH 1965152. indicus Sowerby, Unio : 1866, Conch. Iconica, 16, Unio, pi. 40, species 222 (India, Cuming colln.). Figured holotype BMNH 1965195 ; paratype BMNH 1965196. inflata Orbigny, Castalia : 1835, Mag. de Zool., p. 43 (var. a, rotunda, provincia Corrientesensi, republica Argentina ; var. b, elongata, provincia Chiquitensi, republica Boliviana). Is Castalia ambigua Lamarck, teste, Orbigny, 1846, Voy. Amer. Merid., 5, pt. 3, p. 598, pi. 72, figs. 4-7; 2 syntypes BMNH 1854.12.4.854, both figured. inornattis Reeve, Unio : 1865, Conch. Iconica, 16, Unio, pi. 29, species 147 (Cambojia, Cuming and Hanley collns.). Holotype BMNH [not catalogued] ; specimens from Hanley in ANSP 41673. Non Unio inornatus Lea 1856, changed to Unio verus by Lea, 1870, Synopsis Unio, p. 46. introrugatus Connolly, Unio : 1931, Ann. and Mag. Nat. Hist., (10) 8 : 320, pi. 12, figs. 6-9 (Uganda : Victoria Nyanza). Holotype BMNH 1930.12.3.362 ; paratype BMNH 1930.12.3. 363 and 1937.12.30.1327. involutus "Benson" Hanley, Unio : 1856, Cat. Bivalve Shells, p. 385, pi. 23. fig. 17 (Assam, Hanley colln.). Figured holotype BMNH 1968656. iridella Pilsbry and Frierson, Lampsilis : 1907, Nautilus, 21, pi. 12, two lower right hand figs., 1908, Ibid. 22 : 81 (Valles, [San Louis Potosi Prov.] Mexico). Holotype ANSP 938ioa; paratype BMNH 1908.12.17.5. irisans Marshall, Anodontites : 1926, Proc. United States Natl. Mus., 69 : 10, pi. 2, fig. 35 ; pi. 3, fig. 7 (Venezuela). Holotype USNM 359920 ; 2 paratypes BMNH 1926.2.3.29-30. japanensis Lea, Unio : 1859, Proc. Acad. Nat. Sci. Phila., 11 : 153 (Japan, Cuming colln.); Lea, 1860, Jour. Acad. Nat. Sci. Phila. (2) 4 : 244, pi. 36, fig. 123 ; Lea, 1860, Obs. Unio, 7 : 62. Holotype BMNH 1965181. javona Sowerby, Anodon : 1867, Conch. Iconica, 17, Anodon, pi. n, species 33 (Japan [error for Java fide original label] from Von dem Busch ; Cuming colln.). 3 syntypes BMNH 1965213, none agree exactly with the figure. johnstoni E. A. Smith, Unio (Metaptera) ; 1893, Proc. Zool. Soc. London, p. 640, pi. 59, figs. 18-20 (Lake Mweru, British Central Africa). Figured holotype BMNH 93.8.23.100 ; paratype BMNH 93.8.23.101. jourdyi Morlet [sic], Anodonta : 1886, Jour, de Conch., 34 : 76, pi. 15, fig. i, la (Tonkin [North Vietnam] Environs de Lang-son, Chu). Syntype BMNH 1893.2.4.1395. kelletii Sowerby, Anodon : 1867, Conch. Iconica, 17, Anodon, pi. 19, species 71 (Hab. ?, Cuming colln.). Holotype BMNH 1964392. kirkii Lea, Unio : 1864, Proc. Acad. Nat. Sci. Phila., 16 : 108 (Lake Nyassa, Central Africa, John Kirk) ; Lea, 1866, Jour. Acad. Nat. Sci. Phila. (2) 6 : 32, pi. 12, fig. 30 ; Lea, 1867, Obs. Unio, 11 : 36. Holotype USNM 84056 ; 2 separate valves BMNH 1965154, ex. Cuming colln ; 2 specimens from Kirk's lot, but not seen by Lea, BMNH 62.9.25.7. laevis Haas, Ptychorhynchus : 1910, Ann. and Mag. Nat. Hist. (8) 6 : 498 (Saghalim Island [Siberia]). Holotype Senckenberg Mus. 3626 ; 2 paratypes BMNH 1912.8.16.127-8; all ex Sowerby and Fulton. lampreyanus Baird and Adams, Unio : 1867, Proc. Zool. Soc. London, p. 491, pi. 26, fig. 2 Shanghai, North China). Figured holotype BMNH 67.5.13.16, also refigured by Sowerby 1868, Conch. Iconica, 16, Unio, pi. 87, fig. 469, consists of one valve. lananensis Frierson, Quadrula : 1901, Nautilus, 15 : 75, pi. 4. (Lanana and Banita Creeks near Nacogdoches [Nacogdoches Co.] Texas). Figured holotype ANSP 8 1561 a ; 2 paratypes BMNH 1905.8.15.28-29. lanceolata Lea, Triquetra : 1856, Proc. Acad. Nat. Sci. Phila., 8: 79 (China?). Measured holotype USNM 83884. Changed by Lea to Triquetra contorta, 1856, Proc. Acad. Nat. Sci. Phila., 8 : 301 ; Lea, 1857, Jour. Acad. Nat. Sci. Phila. (2) 3: 319, pi. 33, fig. 3 ; figured UNIONACEA: TYPES AND FIGURED SPECIMENS 87 specimen BMNH 1964400 Shanghai [China], Cuming colln. Lea, 1857, Obs. Unio, 6: 39. Refigured by Sowerby, 1869, Conch. Iconica, 17, Hyria, pi. i, species 2, figs, a— b. laosensis Lea, Unio : 1863, Proc. Acad. Nat. Sci. Phila., 15 : 190 (Laos Mountains, Cambodia, Siam, from Mouhot in Cuming colln.) ; Lea, 1866, Jour Acad, Nat. Sci. Phila. (2) 6 : 63, pi. 21, fig. 6 1 ; Lea, 1867, Obs. Unio, 1 1 : 67. Holotype USNM 86160. A supposed paratype figured by Sowerby, 1866, Conch. Iconica, 16, Unio, pi., 47 species 256, BMNH 1964353, this lot also contains two other specimens which were not seen by Lea, who had but two specimens before him. latialata Sowerby, Hyria : 1869, Conch. Iconica, 17, Hyria, pi. 2, fig. 4 (British Guayana). Holotype BMNH 41.4.29.92 ; paratypes BMNH 41.3.6.30 and 41.3.6.33. layardii Lea, Unio : 1859, Proc. Acad. Nat. Sci. Phila., 11 : 153 (Ceylon, from F. Layard, in Cuming colln.) ; Lea, 1860, Jour. Acad. Nat. Sci. Phila. (2) 4 : 243, pi. 36, fig. 122 ; Lea, 1861 Obs. Unio, 7 : 61. Holotype BMNH 1965159. This was the only specimen seen by Lea. legumen Martens, Anodonta : 1888, Sitzber. Gesell. Natur. Freunde, Berlin, p. 65 (Banhados de S. Leopoddo, Prov. Rio Grande do Sul [Brazil]). Syntype BMNH 91.4.13.56. limnoica Orbigny, Anodonta : 1835, Mag. de Zool., p. 40 (Lagunis, Provincia Corrientesensi, republica Argentina) ; 1846, Voy. Am6r. Merid., 5, pt. 3, pi. 79, figs. 1-3. Holotype BMNH 1854.9.4.26 ; 4 paratypes BMNH 1854.9.4.26/1—4 all labeled, " Mayloya Corrientes. " linguaeformis Morelet, Anodonta : 1875, Series Conch., pt. 4, p. 329, pi. 14, fig. 5 (Cambodje, probablement dans les mare'cages voisins de Battambang). Holotype BMNH 93.2.4.612 ; 3 paratypes BMNH 93.5.12.19-21. luapulaensis Preston, Unio (Nodularia) : 1913, Rev. Zool. Africaine, 3 : 60, pi. 4, fig, n (Confluence of the Lukulu and Luapula Rivers, Belgian Congo). 2 paratypes BMNH 1913.7. 30.24. lucasii Morelet, Anodon : 1851, Jour, de Conch., 2 : 359 (les mare'cages boises de La Calle [Algeria]). Measured holotype BMNH 93.2.4.1950 PL i. fig. 2. ; paratype [not found] figured by Sowerby, 1867, Conch. Iconica, 17, Anodon, pi. 17, species 64, sent to Cuming by Morelet. This appears to be a description of the shell figured, but not described, by Deshayes, 1848, Explor. Sci. de 1'Algerie, Zool., 1, pi. 108, figs. 1-2. If so, Deshayes' named figure is a sufficient indication to afford him priority. lucida Orbigny, Anodonta : 1836, Mag. de Zool., p. 40 (Banda orientali, republica Uruguayensi oriental!) ; 1846, Voy. Amer. Me'rid., 5, pt. 3, p. 620, pi. 79, figs. 4-6 (la riviere dite Canelon grande, Uruguay). Holotype BMNH 1854.12.4.834 ; 3 paratypes BMNH 1854.12.4.834/1-3 from the same lot labeled, " Rio del Rosario, Banda Orientali. " lukuluensis Preston, Mutela : 1913, Rev. Zool. Africaine, 3 : 61, pi. 6, fig. 4 (Confluence of the Lukulu and Luapula Rivers, Belgian Congo). Paratype BMNH 1913.7.30.16. lurulentus Morelet, Anodon : 1849, Testacea Novissima, 1 : 28 (in paludibus fluvii Usumasinta [Peten Prov., Guatemala]). Measured holotype BMNH 93.2.4.2003, figured by Fischer and Crosse, 1894, Miss Sci. au Mexique, pt. 7, 2 : 523, pi. 64, fig. 6 ; 2 paratypes BMNH 93.2.4. 2004-5. The original label reads, " Marais de San Geronimo " [Yucatan, Mexico]. macilenta Morelet, Anodonta : 1845, Moll, de Portugal, p. 102, pi. n (a une lieue de Coimbre, dans de profonds mare'cages voisins du Mondego et connus sous le nom de Val de Geria [Portugal]). Syntype BMNH 93.2.4.1678. mainwairingi " Nevill" Preston, Unio : 1912, Records Indian Mus., Calcutta, 7 : 306 (Siliguri ; also Namtsik, Diha[o]ng [River]). Holotype Indian Mus., figured by Preston 1915, Fauna of British India, Pelecypoda, p. 191, fig. 24 ; 5 paratypes BMNH 91.3.6.3-7. mandarinus Morelet, Unio : 1864, Jour, de Conch., 12 : 159 (Cochinchina [South Vietnam]) ; 3 syntypes BMNH 93.2.4.1971-3 ; Morelet, 1875, Series Conch., pt. 4, p. 354, pi. 17, fig. 2. Listed as a synonym of 17. scobinatus Lea. mandelayensis Theobald, Unio : 1874, Jour. Asiatic Soc. Bengal, 42 (2) : 208, pi. 17, fig. 2 (prope Mandelay, regno Birmanico). Figured holotype BMNH 1888.12.4.2018 ; paratype BMNH 1888.12.4.2019, figured by Hanley and Theobald, 1876, Conch. Indica, p. 62, pi. 154, fig. 4. 88 R. I. JOHNSON rnarocana Pallary, Margaritana : 1918, Bull. Soc. Hist. Nat. Afrique du Nord. 9 : 152 (1'oued Fes [Maroc]). 2 syntypes BMNH 1937.12.30.13034-13035. marteli Pallary, Unio : 1918, Bull. Soc. Hist. Nat. Afrique du Nord, 9 : 151 (La Makina a Fes [Maroc]). 2 syntypes BMNH 1937.12.30.13036-13037. martensi Ihering, Castalina : 1893, Archiv fur Natur., 59 (i) 8i,pl.3, fig. 5 (Rio Camaquam, Rio Grande do Sul, Brazil). Lectotype Senckenberg Mus. 3785, selected by Haas, 1931, Senckenbergiana 13: 41; 2 paralectotypes BMNH 91. 4. 13. 57-58. mashonae Preston, Unio : 1910, Ann. and Mag. Nat. Hist. (8) 6 : 61, pi. 4, fig. 10 (a sluit about 1 6 miles from Eukeldoorn, Mashonaland [South Rhodesia]). Holotype BMNH 1910.1.15.6 ; paratypes BMNH 1910.1.15.7-12. massini Morelet, Unio : 1864, Jour de Conch., 12 : 288 (Cochinchina [South Vietnam]). Holotype BMNH 93.2.4.1590, figured by Morelet, 1875, Series Conch., pt. 4, p. 348, pi. 15, figs, i, 3 ; 2 paratypes BMNH 93.2.4.1591-2. matoniana Orbigny, Unio : 1835, Mag. de Zool., p. 35 (Rio de la Plata, republica Argentina). Is Unio variabilis Wood, teste Orbigny, 1846, Voy. Amer. Merid., 5, pt. 3, p. 604, pi. 71, figs. 1-3. Holotype BMNH 1854.9.4.50 ; paratypes BMNH 1854.9.4.50/1-9. mauritianus Lea, Unio : 1859. Proc. Acad. Nat. Sci. Phila., 11 : 152 (Island of Mauritius, Cuming colln.) ; Lea, 1860, Jour. Acad. Nat. Sci. Phila. (2) 4 : 257, pi. 40, fig. 138 ; Lea, 1860, Obs. Unio, 7 : 75. Holotype BMNH 1965157. melleus Lea, Unio : 1859, Proc. Acad. Nat. Sci. Phila., 11 : 152 (Hab.l [Mexico or Central America], Cuming colln.) ; Lea, 1860, Jour. Acad. Nat. Sci. Phila. (2) 4 : 250, pi. 38, fig. 129 ; Lea, 1860, Obs. Unio, 7 : 68 ; Reeve, 1865, Conch. Iconica, 16, Unio, pi. 21, species 92. Holotype BMNH 1965175 ; also a smaller specimen BMNH 1965176, not seen by Lea. menzieianus "Gray" Reeve, Unio : 1865, Conch. Iconica, 16, Unio, pi. 29, species 152 (New Zealand, Cuming colln.). BMNH [lost] [is Unio tnenziesi Gray], menziesi Gray, Unio : 1843, [in] Dieffenbach, Travels in New Zealand 2 : 257 (Lake Taupo, North Island, New Zealand, restricted by Dell, 1953, Trans. Royal Soc. New Zealand, 81 : 229). Holotype BMNH 42.5.17.170, figured by McMichael and Hiscock, 1958, Australian Jour. Marine and F. W. Res., 9, pi. 14, figs. 1-2. merdiger Reeve, Unio : Conch. Iconica, 16, Unio, pi. 28, species 145 (Hungary, Cuming colln.). Holotype BMNH 1964161. mexicanus Sowerby, Unio : 1867, Conch. Iconica, 16, Unio, pi. 55, species 281 (Mexico, Taylor colln.). Holotype BMNH 79.2.26.249. micans Anthony, Anodon : 1865, Amer. Jour. Conch., 1 : 162, pi. 16, fig. i (Texas). Holo- type MCZ 187290 ; paratype BMNH 1968652 figured by Sowerby 1867, Conch. Iconica, 17, Anodon, pi. 15, species 53. micropterus Morelet, Unio : 1866, Jour, de Conch., 14 : 63 (in torrentibus montanis Cam- bodiae). Measured holotype BMNH 93.2.4.1596, figured by Morelet, 1875, Series Conch., pt. 4, p. 349, pi. 15, fig. 6 (the figure is reduced) ; paratype BMNH 93.2.4.1597. minuana Orbigny, Monocondylaea : 1835, Mag. de Zool., p. 37 (Uruguayensis orientali republica) ; 1846, Voy. Amer. Merid., 5, pt. 3, p. 612, pi. 70, figs. 8-10 (dans les rivieres de Canelon Grande et del Rosario, dans la Banda oriental do la Plata, Uruguay). Holotype BMNH 1854.9.4.17 ; 2 paratypes BMNH 1854.9.4.27/1-2. misellus Morelet, Unio : 1865, Jour, de Conch. 13 : 21 ([Salaburi] Siam). Holotype BMNH 93.2.4.1593, figured by Morelet, 1875, Series Conch., pt. 4, p. 341, pi. 14, fig. 2 ; 2 paratypes BMNH 93.2.4.1994-5. moretonicus Sowerby, Unio : 1865, Conch. Iconica, 16, Unio, pi. 24, species 118 (Moreton Bay, Australia). Holotype BMNH 1958.5.16.1. morini Morelet, Unio : 1851, Testacea Novissima, 2 : 25 (flumen Usumasinta [Peten Prov., Guatemala]). Measured holotype BMNH 93.2.4.2026, figured by Fischer and Crosse, 1894, Miss. Sci. au Mexique, pt. 7, 2 : 576, pi. 60, fig. 2, pi. 67, fig. 4. mouhoft Lea, Monocondylaea : 1863, Proc. Acad. Nat. Sci, Phila., 15 : 190 (Laos Mountains, Cambodia, Siam) ; changed without explanation to : mouhotiana Lea, Monocondylaea : 1866, Jour. Acad. Nat. Sci. Phila. (2) 6 : 65, pi. 21, fig. UNIONACEA: TYPES AND FIGURED SPECIMENS 89 62 ; Lea, 1867, Obs. Unio, 11 : 69. Holotype USNM 86339 ; paratype BMNH [not cata- logued]. tnucidus Morelet, Unio : 1845, Moll, de Portugal, p. in, pi. 14, fig. 3 (La Tamega, le Cavado et la Lima [Rivers, Portugal]). Holotype BMNH 93.2.4.1974 ; paratype BMNH 93.2.4.1975, both from le Cavado. The holotype is the measured specimen. The figure appears to be a composite of both specimens; 2 additional paratypes BMNH 93.2.4.1979-80 from Rio Tamega and Mondago. mutabilis Lea, Unio : 1859, Proc. Acad. Nat. Sci. Phila., 11 : 152 (Brisbane water, Australia ; New Zealand, H. Cuming ; Murray River, Australia, W. Newcomb) ; Lea, 1860 Jour. Acad. Nat. Sci. Phila. (2) 4 : 248, pi. 38, fig. 127 ; Lea, 1860, Obs. Unio, 7 : 60. Figured holotype BMNH [lost] from New Zealand. Lectotype, here selected, BMNH 1965153 from Brisbane water, figured by Reeve, 1865, Conch. Iconica, 16, Unio, pi. 24, species 1 12. It was refigured by McMichael and Hiscock, 1958, Australian Jour. Mar. F. W. Res., 9 : 449, pi. n, fig. n ; paralectotype USNM 85815, Ibid., pi. n, figs. 8-9 ; paralectotype USNM 84405, from Murray River. mweruensis E. A. Smith, Unio : 1908, Proc. Mai. Soc. London, 8 : 13, text figs. (Lake Mweru, Central Africa). Figured syntype BMNH 1907.11.11.34 ; also syntypes BMNH 1907. II-33-35-49- navigioliformis Lea, Unio : 1859, Proc. Acad. Nat. Sci. Phila., 11 : 152 (Hab. ?, Cuming colln.) ; Lea, 1860, Jour. Acad. Nat. Sci. Phila. (2) 4 : 248, pi. 37, fig. 124 ; Lea, 1860, Obs. Unio, 7 : 63. Holotype, and only specimen seen by Lea, not found in the BMNH or in the USNM. nehringi Ihering, Castalina : 1893, Archiv. fur Natur., 59 (i) : 75, pi. 3, fig. 4 (Rio Piracicabo, Sao Paulo, Brazil). Lectotype Senckenberg Mus. 3787, selected by Haas, 1931, Senckenbergiana 13: 41; paralectotype BMNH 1891.4.13.116. nehringi Ihering, Glabaris : 1893, Archiv fur Natur.; 59 ( i) : 60 (Rio St. Maria, Rio Grande do Sul ; [Rio] Piracicaba, Sao Paulo, Brazil). 3 paratypes BMNH 1891.4.13.107-109. nopalatensis Sowerby, Anodon : 1867, Conch. Iconica, 17, Anodon, pi. 16, species 58 (River Nopalata, Mexico). Holotype BMNH 1965189, consists of opposite of figured valve only. novaehollandiae Gray, Unio : 1834, Proc. Zool. Soc. London, pt. 2, p. 57 (in novae Hollandiae flumine Macquarrier [probably Port Macquarie, Hastings River, N. coast of New South Wales]). Syntypes BMNH 1880.4.10.6-7, consisting of 2 left valves, figured by McMichael and Hiscock 1958, Australian Jour. Mar. and F. W. Res., 9 : 474, pi. 15, figs. 1-2. nyassae Sowerby, Unio : 1866, Conch. Iconica, 16, Unio, pi. 41, species 224, figs, a, b (Lake Nyassa, [Central Africa] Cuming colln.). BMNH [lost]. nyassaensis Lea, Spatha : 1864, Proc. Acad. Nat. Sci. Phila., 16 : 109 (Lake Nyassa, Central Africa, John Kirk) ; Lea, 1866, Jour. Acad. Nat. Sci. Phila. (2) 6 : 36, pi. 13, fig. 33 ; Lea, 1867, Obs. Unio, 11,: 40. Holotype USNM 86777 '• specimen from Kirk's lot, but not seen by Lea, BMNH 1864.5.14.1. nyassaensis Lea, Unio : 1864, Proc. Acad. Nat. Sci. Phila., 16 : 108 (Lake Nyassa, Central Africa, John Kirk) ; Lea, 1866, Jour. Acad. Nat. Sci. Phila. (2) 6 : 33, pi. 12, fig. 32 ; Lea, 1867, Obs. Unio, 11 : 40. Holotype USNM 84057 ; specimen from Kirk's lot but not seen by Lea, BMNH 1864.5.14.3. obesa Hanley and Theobald, Unio marginalis : 1872, Conch. Indica, p. 20. pi. 43, fig. 3 (River Irawadi, Birmah). Figured holotype BNMH 1907.12.30.46 ; idiotype BMNH 88.12.4.1642, from Tonyhu, Pegu ex Theobald. obicularis Morelet, Monocondylus : 1866, Rev. et Mag. de Zool. (2) 18 : 167 (no locality [Battambang, Cambodje]). Measured holotype BMNH 93.2.4.1982, figured by Morelet, 1875, Series Conch., pt. 4, p. 338, pi. 16, fig. 5, the figure is slightly enlarged. obliqua Longstaff, Nodularia (Caelatura) parreysii : 1914, Jour. Linn. Soc. London, 32 : 255, pi. 1 8, fig. ii (White Nile River at Tawili, Masran Island and Melut, Southern Sudan). Figured holotype BMNH 1923.6.8.1214, from Melut. obliquiradiatus Reeve, Unio : 1865, Conch. Iconica, 16, Unio, pi. 29, species 151 (Hab. ?, Cuming colln.). BMNH [lost]. 90 R. I. JOHNSON obliterata Fischer and Crosse, Unio scutulatus : see under scutulatus Morelet, Unio. oleivorus Heude, Mycetopus : 1877, Conch. Fluv. Nanking, pt. 3, pi. 22, fig. 46 ; pi. 23, fig. 48 (La Hoai superieure, department de Ing-tch'eou [China]). BMNH 99.4.22.69, labeled Ngan-hou6, from R. P. Heude in 1878, ex A. Morelet colln. Not a primary type. ortmanni Frierson, Unio : 1913, Nautilus, 27 : 14, pi. 2 (Conchins River, near Quirigua, Guatemala [Atlantic Drainage]). Figured holotype ANSP i77544a ; 3 paratypes BMNH 1914.1.7.305-307. ostreatus Morelet, Unio : 1849, Testacea Novissima, 1 : 29 (flumen Usumasinta [Peten Prov., Guatemala]). Measured holotype BMNH 94.3.22.2, figured by Fischer and Crosse, 1894, Miss. Sci. au Mexique, pt. 7, 2 : 573, pi. 63. fig. 3, pi. 70, fig. 4. ovata Sowerby, Castalia : 1869, Conch. Iconica, 17, pi. i, species 4 (Brazil, Orbigny colln.). Holotype BMNH 1964402. ozarkensis Call, Unio : 1887, Proc. United States Natl. Mus., 10 : 498, pi. 27 (Currant River, Shannon Co., Missouri, and in Jack's Fork and Big Creek, tributaries of it). Holotype MCZ 5707 ; paratype BMNH 98.2.1.41, localities not separated. pahangensis E. A. Smith, Unio : 1899, Proc. Mai. Soc. London, 3 : 315, text fig. (Pahang River, Malay Peninsula). Figured holotype BMNH 99.9.4.1. paivaeanus Morelet, Unio : 1865, Jour, de Conch., 13 : 227 (Siam [riviere Saraburi]). Holo- type BMNH 93.2.4.1738, figured by Morelet, 1875, Series Conch., pt. 4, p. 353, pi. 17, fig. 7, where it is listed as a synonym of U. rusticus Lea. pajakomboensis Bullen, Unio : 1906, Proc. Mai. Soc. London, 7 : 15, pi. 2, figs. 9-11 (River at Pajakombo, Sumatra). Figured holotype BMNH 1906.1.16.52. pallaryi Longstaff, Nodularia (Lanceolaria) teretiuscula : 1914, Jour. Linn. Soc., London, 32 : 256, pi. 18, figs. 12-14 (White Nile River, Mogran, Southern Sudan). Figured holotype BMNH 1923.6.8.1214. pallegoixi Sowerby, Anodon : 1865, Conch. Iconica, 17, Anodon, pi. 8, species 18, fig. 17 (Siam, Cuming colln.). Holotype BMNH 1965193 ; paratype BMNH 1965194. pallida Anthony, Anodon : 1865, Amer. Jour. Conch., 1 : 162, pi. 15, fig. 3 (Michigan). Holo- type MCZ 161871 ; 3 paratypes BMNH 1965155. paludosus Morelet, Unio : 1849, Testacea Novissma, 1 : 30 (paludosa prope San Geronimo Yucataneorum [Mexico]). Measured holotype BMNH 93.2.4.2024, figured by Fischer and Crosse 1894, Miss. Sci. au Mexique, pt. 7, 2 : 559, pi. 59, fig. 3. panacoensis Von dem Busch, Unio : 1843 [in] Philippi, Abb. und Besch. Conch., 1 : 75, pi. 2 (flumen Panaco prope Tampico [Mexico]). Paratype BMNH 1965183. papyracea Anthony, Anodon : 1865, Amer. Jour. Conch., 1 : 161, pi. 15, fig. 2 (Hab. ? [Poto- mac River, Virginia]). Holotype MCZ 150656 ; 3 paratypes BMNH 1965151. paraguayana Orbigny, Monocondylaea : 1835, Mag. de Zool., p. 37 (Rio Parana [pres du village d'ltaty] et Rio Batel, provincia Corrientesensi, republica Argentina) ; 1846, Voy. Amer. Merid., 5, pt. 3, p. 612, pi. 70, figs. 5-7. Holotype BMNH 1854.9.4.13 ; 4 paratypes 1854.9.4.13/1-4. parchappii Orbigny, Monocondylaea : 1835, Mag. de Zool., p. 38 (Rio Parana, provincia Corrientesensi, republica Argentina) ; 1846, Voy. Amer. Merid., 5, pt. 3, p. 613, pi. 68, figs. 1-3 (sous les pierres de la cote du Parana, aux environs d'ltaty, province de Corrientes, Argentine). Holotype BMNH 54.9.4.29 ; 7 paratypes BMNH 1854.9.4.29/1-7. parma "Benson" Sowerby, Unio : 1868, Conch. Iconica, 16, Unio, pi. 95, species 514 (Tennas- serim, East Indies, Hanley colln). Figured holotype BMNH 88.12.4.1669 ; refigured by Hanley and Theobald 1876, Conch. Indica, p. 61, pi. 154, fig. i. patagonica Orbigny, Unio : 1835, Mag. de Zool., p. 37 (Rio Negro, Patagonia) ; 1846, Voy. Amer. Merid., 5, pt. 3, p. 610, pi. 70, figs. 1-4. Figured holotype BMNH 1854.12.4.850 ; 7 paratypes BMNH 1854.12.4.850/1-7 ; 5 smaller paratypes BMNH 1854.12.4.851. pellis-lacerti Morelet, Unio : 1865, Jour, de Conch., 13 : 27 (Siam [dans le riviere de Saraburi ; Cambodje, dans celle de Battambang, et en Cochinchine, pres de Mitho]). Holotype BMNH 93.2.4.1969, figured by Morelet, 1875, Series Conch., pt. 4, p. 356, pi. 17, fig. 5, paratypes BMNH 93.2.4.1966-8 and 1970, all from the first locality. UNIONACEA: TYPES AND FIGURED SPECIMENS 91 persulcatus Lea, Unio : 1859, Proc. Acad. Nat. Sci. Phila., 11 : 153 (Mexico, Hanley colln.) ; Lea, 1860, Jour. Acad. Nat. Sci. Phila. (2) 4 ; 255, pi. 40, fig. 135 ; Lea, 1860 Obs. Unio, 7 : 73. Holotype BMNH 1900.3.19.22. pinei Wright, Unio : 1897, Nautilus, 11 : 40 (unnamed lake in Witthacoochee [Withlacoochee] River region of Hernando Co., Florida). Holotype USNM 150127, figured by Simpson 1900, Proc. Acad. Nat. Sci. Phila., 52 : 80, pi. 3, fig. i ; refigured by Johnson 1967, Occ. Papers on Moll., 3 : 8, pi. 10, fig. 5 ; 2 paratypes BMNH 1905. 8.15.34-35. piracicabana Ihering, Unio aethiops : 1893, Archiv. fiirNatur., 59 (2) : 102 ([Rio] Piricacaba, Sao Paulo, Brazil). Holotype Senkenberg Mus. 4031 ; 2 paratypes BMNH 91.4.13.119-120. planivalvis Morelet, Unio : 1851, Testacea Novissima, 2 : 24 (in paludibus fluminis Usuma- sinta vicinis [Peten Prov., Guatemala]). Measured holotype BMNH 93.2.4.2025, figured by Fischer and Crosse, 1894, Miss. Sci. au Mexique, pt. 7, 2 : 560, pi. 59, fig. 2. The original label reads, " Marais de S. Geronimo " [Yucatan, Mexico]. plexoides Frierson, Psoronaias : 1927, Check List North American Naiades, p. 63. Based on Sowerby, 1868, Conch. Iconica, 16, Unio, pi. 64, species 324 (Siam, Taylor colln.). Not known if ever in BMNH. plicatulus Lea, Unio : 1859, Proc. Acad. Nat. Sci. Phila., 11 : 152 (Borneo, Cuming colln.) ; Lea, 1860, Jour. Acad. Nat. Sci. Phila. (2) 4 : 247, pi. 37, fig. 126 ; Lea, 1860, Obs. Unio, 7 : 65. Holotype BMNH 1965156. plicatus Leach, Dipas ; 1814, Zool. Miscellany, 1 : 120, pi. 55 (Hab. ?, British Museum). Holotype BMNH 1952.5.10.1 from the Hans Sloane colln., consisting of two unmatched valves, refigured by Wilkins, 1953, Bull. Brit. Mus. Nat. Hist., Hist. Ser., 1 : 40, pi. 12. plicatus Sowerby, Mycetopus : 1868, Conch. Iconica, 16, Mycetopus, pi. 2, species 3 (Hab. ?). Holotype BMNH 46.7.24.18 ; paratype BMNH 46.7.24.19. pliciferus Reeve, Unio : 1864, Conch. Iconica, 16, Unio, pi. 10, species 37 (Mexico, Cuming colln.). BMNH [lost]. pliculosus Martens Unio : 1894, Sitzber. Gesell. Natur. Freunde, Berlin, p. 216 (Singei [Prov. Hwanghaido] (a) und zwischen Okkwa und Chhangpyong (b) ersteres im siidlichen letzteres im nordlichen Theil von Korea) ; Martens, 1905, Zool. Jahrb., Suppl. 8, p. 61, pi. 3, figs. 3, 3b. Paratype BMNH 94.11.24.8, from locality (b). praestricta Fischer and Crosse, Unio crocodilorum : see under crocodilorum Morelet, Unio. pressiorstris Martens, Unio : 1900, Natur. Deutschen Mai. Ges., 32 : 14 (Lake Danau-Baru, Indragiri, Sumatra). Measured holotype figured by Haas 1914, [in] Martini and Chemnitz, Conch. Cabinet (2) 9, pt. 2, sec. 2, p. 227, pi. 27, fig. i ; paratype BMNH 1901.6.14.4. prolongata Fischer and Crosse, Unio calamitarum : 1894, Miss. Sci. au Mexique, pt. 7, 2 : 613, pi. 63, figs. 5, 5a ([Mexico] Morelet colln.). Figured holotype BMNH 93.2.4.2009. psammoica Orbigny, Unio : 1835, Mag. de Zool., p. 35 (Rio Parana, provincia Corrientesensi, republica Argentina) ; 1846, Voy. Amer. Merid., 5, pt. 3, p. 608, pi. 71, figs. 4-7 (la Parana, pres du village d'ltaty, bien au-dessus de Corrientes). Lectotype, here selected BMNH 54.12.4.849, which is the specimen figured on pi. 71, figs. 4-6 ; 2 smaller paralectotypes BMNH 1854.4.12.849/1-2, the smallest specimen is figured on pi. 71, fig. 7. psoricus Morelet, Unio : 1851, Testacea Novissima, 2 : 25 (flumen Usumasinta [Peten Prov., Guatemala]). Measured holotype BMNH 93.2.4.2033, figured by Fischer and Crosse, 1894, Miss. Sci. au Mexique, pt. 7, 2 : 572, pi. 61, fig. 3 ; paratype BMNH 93.2.4.2034. puelchana Orbigny, Anodonta : 1835, Mag. de Zool., p. 40 (Rio Negro, Patagonia) ; 1846, Voy. Am£r. Me"rid., 5, pt. 3, p. 620, pi. 79, figs. 7-9 (les marais de San-Xavier, a six lieues au- dessus du Carmen, sur le Rio Negro, Patagonie). Holotype BMNH 1854.9.4.21 ; 3 smaller paratypes [under same number]. punctatus Preston, Mycetopus : 1909, Ann. and Mag. Nat. Hist. (8) 3 : 513, pi. 10, fig. 8 (Rio Chemchi, U.S. Columbia). Figured holotype BMNH 1915.1.6.83. quadrata Sowerby, Castalia : 1869, Conch. Iconica, 17, Castalia, pi. 2, species 7, figs, a-b (Hab. ?.). Figured holotype BMNH 49.1.5.5. quadrilatera Orbigny, Castalia : 1835, Mag. de Zool., p. 42 (provincia Moxensi, republica Boliviana) ; 1846, Voy. Am6r. Merid. 5, pt. 3, p. 599, pi. 73 (Rio de San-Miguel, Guarayos, B* 92 R. I. JOHNSON Bolivia). Lectotype, here selected, BMNH 54.9.4.2 the largest specimen figured ; figured paralectotype BMNH 1854.9.4.2/1 ; 6 paralectotypes BMNH 1854.9.4.2/2-7. ranarutn Morelet, Anodonta : 1845, Moll, de Portugal, p. 104, pi. 12, fig. 2 (affluens de la Guadiana qui descendent des hautes-valle"es de 1'Algarve entre Mertola et Castro- Verde [Portu- gal]). Holotype BMNH 93.2.4.1737. ravistellus Morelet, Unio : 1849, Testacea Novissima, 1 : 29 (in lacu Yzabal, republicae Guate- malensis). Measured holotype BMNH 93.2.4.2021, figured by Fischer and Crosse, 1894, Miss. Sci. au Mexique, pt. 7, 2 : 509, pi. 61, fig. 4 ; 2 paratypes BMNH 93.2.4.2022-3. recta Sowerby, Hyria : 1869, Conch. Iconica, 17, Hyria, pi. 5, species 10 (Hab. ?, Cumingcolln.). BMNH [lost]. rectilinearis Sowerby, Unio : 1868, Conch. Iconica, 16, Unio, pi. 65, species 332 (River Colum- bia [South Africa] Taylor colln.). Holotype BMNH 74.12.11.4. regularis Morelet, Anodonta : 1845, Moll, de Portugal, p. 100, pi. 10 (La Tamega . . . aux en- virons de Chaves [Portugal]). Syntype BMNH 93.2.4.1679. reticulatus Sowerby, Anodon : 1867, Conch. Iconica, 17, Anodon, pi. 10, species 27 (River Amazon [Brazil], Cuming colln.). Holotype BMNH 1965208. rhuacoica Orbigny, Unio : 1835, Mag. de Zool., p. 36 (Banda orientali, republica Uruguayensi orientali) ; 1846, Voy. Ame"r. Merid., 5, pt. 3, p. 606, pi. 69, figs. 4-5 ; pi. 71, figs. 12-14 (un ris- seau pres de Maldonado, et dans le Rio Canelon grande pres de Montevideo, Uruguay). Lectotype, here selected, BMNH 1854.12.4.838 specimen figured on pi. 69, figs. 4-5 ; paralectotypes BMNH 1854.12.4.838/1-3 all from the latter locality. riograndensis Ihering, Anodonta : 1890, Archiv. fur Natur., 56 (i) : 154 ([RioCamaquam] Rio Grande do Sul; und dem Plata Gebeite, Brazil). Holotype Senkenberg Mus, 3838 ; 4 paratypes BMNH 91.4.13.15-18, localities not separated. robsoni Frierson, Parreysia : 1927, Check List of North American Naiades, p. 44. (East Indies). Name based on shell figured by Reeve, 1866, Conch. Iconica, 16, Unio, pi. 36, species 198 as Unio pumilis Lea. Holotype BMNH 1965150. rosea Bernardi, Galatea concamerata : 1860, Monog. Galatea et Fischeria, p. 20, pi. 3, figs. 1-2 ([Africa] Cuming colln.). BMNH [lost]. rostratus Sowerby, Anodon : 1867, Conch. Iconica, 17, Anodon, pi. 2, species 4 (Lao, Sowerby colln.). 3 syntypes BMNH 1965214 all smaller than figured specimen. rotundatus Reeve, Unio : 1865, Conch. Iconica, 16, Unio, pi. 23, species 106 (North America, Cuming colln.). BMNH [lost]. rugatus Sowerby, Mycetopus : 1868, Conch. Iconica, 16, Mycetopus, pi. 3, species 7 (Victoria River, Australia). Lectotype BMNH 44.1.12.160, selected by McMichael and Hiscock, 1958, Aust. Jour. Marine and F. W. Res., 9 : 433, pi. 9, figs. 1-2. sacculus "Anthony" Reeve, Unio : 1864, Conch. Iconica, 16, Unio, pi. 15, species 67 (North America, Cuming colln). BMNH [lost]. santa-mariae Simpson, Diplodon : 1914, Descr. Cat. Naiades, 3 : 1270 (Brazil?). Specimens BMNH 1891.4.13.24-26, from Rio St. Maria, Rio Grande do Sul, Brazil, distributed by Ihering but never described by him. scamnatus Morelet, Unio : 1849, Testacea Novissima, 1 : 30 ([Rio Las Pazas] Cacajajicara, [Pinar del Rio] Cuba). Lectotype, here selected, BMNH 1893.2.4.1976, labeled, " Rio Tacotaco, " [Pinar del Rio] Cuba, plate 2 fig. 3. Length 52, height 34, width 17 mm ; paralectotypes BMNH 1893.2.4.1977-78, one of which consists of one valve. schadei Marshall, Anodontites : 1943, Jour. Washington Acad. Sci., 24 : 78 [unnumbered plate], figs. 4-6 (Tubicuary River, Aroja, Paraguay). Holotype USNM 434732 ; paratype USNM 434837, the only specimens seen by Marshall, who received them from Fulton ; 2 additional specimens BMNH 1934.7.3.20-21. schombergiana Sowerby, Castalia : 1869, Conch. Iconica, 17, Castalia, pi. i, species 3 (British Guayana, Schomberg). Holotype BMNH 1841.1.26.5. schomburgianus Sowerby, Anodon : 1870, Conch. Iconica, 17, Anodon, pi. 34, species 137 (British Guiana, Schomberg). BMNH [lost]. scnomburgki Martens, Anodonta (Lamproscapha) : 1860, Proc. Zool. Soc. London, 28 : 15 UNIONACEA: TYPES AND FIGURED SPECIMENS 93 (Siam). Holotype BMNH 59.5.23.8, figured by Haas, 1920, [in] Martini and Chemnitz, Conch. Cab. (2) 9, pt. 2, sec. 2, p. 296, pi. 36, fig. 4. scriptus Sowerby, Anodon : 1867, Conch. Iconica, 17, Anodon, pi. 4, species 9 (Hab. ?, Cuming colln.). Holotype BMNH 1964370. scutulatus Morelet, Unio : 1849, Testacea Novissima, 1 : 30 (propre S. Geronimo, Yucatan- eorum [Mexico]). Measured holotype BMNH 93.2.4.2014, figured by Fischer and Crosse, 1894, Miss. Sci. au Mexique, pt. 7, 2 : 561, pi. 59, fig. 4 ; 2 paratypes BMNH 93.2.4.2017-18 BMNH 1893.2.4.2015-16 were also figured, pi. 59, figs. 5 and 6, and named varieties obliterata and secabilis respectively. scutum Sowerby, Unio : 1868, Conch. Iconica, 16, Unio, pi. 94, species 510 (Tenasserium, Hanley colln.). Holotype BMNH 1907.10.28.239. secabilis Fischer and Crosse, Unio scutulatus : see under scutulatus Morelet, Unio. semicorrugata Preston, Spatha : 1909, Ann. and Mag. Nat. Hist. (8) 4 : 90, pi. 4, fig. 7 (Lower Congo). Paratype BMNH [not catalogued]. semigranosus Reeve, Unio : 1864, Conch. Iconica, 16, Unio, pi. 10, species 36 (Mexico, Cuming colln.). Holotype BMNH 196496. Non Unio semigranosus Von dem Busch 1845. semipustulata Fischer and Crosse, Unio crocodilorum : See under crocodilorum Morelet, Unio. semiquadrata Sowerby, Unio : 1866, Conch. Iconica, 16, Unio, pi. 48, species 258 (Camboja, Lao Mountains, Cuming colln.). BMNH [lost]. semisulcata H. Adams, Monocondylaea (Plagiodon) : 1870, Proc. Zool. Soc. London, p. 376, pi. 27, fig. 3 (East Peru). Syntype BMNH 1907.10.28.128, smaller than figured specimen. senegalensis Lea, Anodonta : 1859, Proc. Acad. Nat. Sci. Phila., 11 : 154 (Senegal, Jay, Verreaux, Cuming) ; Lea, 1860, Jour. Acad. Nat. Sci. Phila. (2) 4 : 260, pi. 41, fig. 140 ; Lea, 1860, Obs. Unio, 7 : 78. Sowerby, 1867, Conch. Iconica, 17, Anodon, pi. n, species 35. Holotype BMNH 1965167 ; paratype BMNH 1965168. senilis Martens, Spatha trapezia : 1897, Bes. Weicht. Deutsch-Ost-Afrikas, p. 244, text fig. (Victoria Nyansa, Nordwestseite, bei Towalio, Buddu Kiiste, und Insel Soweh in Uganda). Paratype BMNH 1902. 5.26.45. from Buddu Kiiste. servainiana Bourguignat, Grandidieria : 1885, Bull. Soc. Mai. de France, 2 : 6. Name based on shell figured as Unio burtoni Smith 1881, Proc. Zool. Soc. London, p. 297, pi. 34, fig. 33, non Woodward 1859. Figured holotype BMNH 80.12.20.21. shambiensis Longstaff, Nodularia (Caelatura) : 1914, Jour. Linn. Soc. London, 32 : 253, pi. 18, figs. 4-7 (Lake ShambS [Bahrel-Gebel] Southern Sudan). 3 paratypes BMNH 1923.6.8. 1214-16. shanghaiensis Lea, Unio : 1859, Proc. Acad. Nat. Sci. Phila., 11 : 153 (Shanghai, China, Cuming) ; Lea, 1860, Jour. Acad. Nat. Sci. Phila. (2) 4 : 242, pi. 36, fig. 121 ; Lea, 1860, Obs. Unio, 7 : 60. Reeve, 1865, Conch. Iconica, 16, Unio, pi. 21, fig. 96. Holotype BMNH 1 965 1 71 ; a smaller specimen not seen by Lea BMNH 1965172. siculus Swainson, Unio : 1840, Treatise on Malacology, p. 282, fig. 58. Hanley, 1856, Cat. Recent Bivalve Shells, p. 383, Supp. pi. 20, fig. 19 (Lake Leontini, Sicily). Probable syntype BMNH 1907.10.28.255, fide Hanley. Probable syntype figured by Sowerby, 1868, Conch. Iconica, 16, Unio, pi. 71, species 255, BMNH 1907.12.30.366 ex Hanley colln. sifefe/menste Lea, Unio : 1859, Proc. Acad. Nat. Sci. Phila., 11 : 151 (Sikkim, India, Cuming and Hanley) ; Lea, 1859, Jour. Acad. Nat. Sci, Phila. (2) 4 : 251, pi. 39, fig. 131; Lea, 1860, Obs. Unio, 7 : 69. Holotype BMNH 1965177 Cuming colln. ; a second specimen not seen by Lea BMNH 1965178 ; paratype BMNH 1900.3.19.23 Hanley colln. siliquosus Orbigny, Mycetopoda : 1835, Mag. de Zool., p. 41 (provincia Corrientesensi, republica Argentina ; Santa-Cruz de la Sierra, republica Boliviana) ; 1846, Voy. Ame'r. Merid., 5, pt. 3, p. 601, pi. 67 (le cours de Parana, province de Corrientes, r^publique Argen- tine ; tous les affluens de 1'Amazone, provinces de Santa-Cruz de Chiquitos et de Moxos, re'publique de Bolivia) [not located]. Non Spix 1827, teste, Ihering 1910, Abh. Senckenb. Ges., 32, p. 121 and named Mycetopoda orbignyi. Paratype BMNH [not located] figured by Sowerby 1868, Conch. Iconica, 16, Mycetopus, pi. 3, species 2, fig. 2a. 94 R- I. JOHNSON sitnonis Tristram, Unto : 1865, Proc. Zool. Soc. London, p. 544 (the Jordan, the sea of Galilee, the Orontes and the Leontes (Litany) [Palestine]). Measured holotype BMNH 1936.3.10.6, plate 2 fig. 2 from the Orontes River. sitnplicidus Sowerby, Anodon : 1867, Conch. Iconica, 17, Anodon, pi. 12, species 40 (Hab. ?, Taylor colln.). Not known if ever in BMNH. singleyanus Marsh, Unio : 1891, Nautilus, 5 : 29, figured by Simpson, Proc. United States Natl. Mus., 15 : 426, pi. 68, figs. 4, 5 (a small creek near Pilatka [Palatka, Putnam Co.] Florida). Idiotype BMNH 98.2.1.42, from Sumter Co., Florida. sirionos Orbigny, Anodonta : 1835, Mag. de Zool., p. 40 (var. Major, provincia Chiquitensi, Boliviana ; var. Minor, provincia Corrientesensi, republica Argentina) ; 1846, Voy. Amer. Merid., 5, pt. 3, p. 615, pi. 74, figs. 4-6 ; pi. 80, figs. 1-4 (le Rio Canelon Grande, pres de Montevideo, du Parana, au-dessus de Corrientes, pres du village d'ltaty, Argentina et dans le Rio de San-Miguel, au pays des Guarayos, province de Chiquitos, et dans le Piray, province de Santa-Cruz en Bolivia). Holotype BMNH 54.9.4.9 ; 6 smaller paratypes BMNH 1854.9.4. 9/1-6 all from le Rio-San-Miguel au pays des Guarayos, Chiquitos ; 3 paratypes BMNH 1854.9.4.35-36. sitifensis Morelet, Unio : 1851, Jour, de Conch., 2 : 360 (1'oued sefsaf, pres de Philippeville ; [Algeria]). Lectotype, here selected, BMNH 93.2.4.1965, plate 2 fig. 6, labeled Algerie, " 1'oued dehhab, pres de Bone [sic] ". Length 71, height 33, width 23 mm. srnithi Bourguignat, Grandidieria : 1885, Bull. Soc. Mai. de France, 2 : 7. Name based on shell figured as Unio burtoni Smith 1881, Proc. Zool. Soc. London, p. 297, pi. 34, fig. 33a, non Woodward 1859. Figured holotype BMNH 80.12.20.31. sobaensis Preston, Nodularia : 1914, Jour. Linn. Soc. London, 32 : 266, pi. 18, figs. 1-3 (Soba, Blue Nile). Figured holotype BMNH 1923.6.8.1210 ; 2 paratypes BMNH 1923.6.8.1211-1212. solenidea Sowerby, Anodon : 1867, Conch. Iconica, 17, Anodon, pi. 18. species 65 (Rio Francisco [Brazil] Cuming colln.). Holotype BMNH 1965201 ; paratype BMNH 1965202. soleniformis Orbigny, Anodonta : 1835, Mag. de Zool., p. 41 (no locality) ; 1846, Voy. Amer. Merid., 5, pt. 3, p. 617, pi. 74, figs. 1-3 (Iribucua, dans le Parana, bien audessus de Corrientes ; et dans le Rio Balel [Argentina]). Holotype BMNH 1854.12.4.857 and paratypes BMNH 1854.12.4.857/1-10, from the first locality. soleniformis Orbigny, Mycetopus : 1835, Mag. de Zool., p. 41 (Santa-Cruz de la Sierra, rep. Boliviana) ; 1846, Voy. Am6r. Merid., 5, pt. 3, p. 601, pi. 66 (Rio Piray, entre Santa-Cruz de la Sierra et la province de Moxos, Bolivia). Holotype BMNH 1854.12.4.859, the figure is slightly reduced ; paratypes BMNH 1854.12.4.859. solisiana Orbigny, Unio : 1835, Mag. de Zool., p. 36 (Rio de la Plata, in provincia Buenos-Ayres republica Argentina) ; 1846, Voy. AmeY. Me"rid., 5, pt. 3, p. 604, pi. 69, figs. 1-3 (plages sablon- neuses de la Plata, aux environs de Buenos-Ayres ; elle y est tres-commune, surtout a 1'em- bouchure des petites ruiaux, de ceux de Punta Lara et de Maldonado [Argentina]). Holotype BMNH 1854.9.4.47 ; ii paratypes BMNH 1854.9.4.48, localities not separated. spekii, Woodward, Iridina (Pleiodon) : 1859, Proc. Zool. Soc. London, 27 : 348, pi. 27, fig. 2 (Lake Tanganyika [Central Africa]). Holotype BMNH 1859.12.23.8, refigured by Sowerby 1866, Conch. Iconica, 16, Pleiodon, pi. i, species 2. spheniopsis Morelet, Unio : 1849, Testacea Novissima, 1 : 29 (regiones superiores fluminis Usumasinta [Penten Prov., Guatemala]). Measured holotype BMNH 93.2.4.2028, figured by Fischer and Crosse, 1894, Miss. Sci. au Mexique, pt. 7, 2 : 583, pi. 61, figs. 2, 2a, 2b ; smaller paratype BMNH 93.2.4.2029. spixii Orbigny, Anodonta : 1835, Mag. de Zool., p. 39 (Rio-Parana, provincia Corrientesensi, republica Argentina). New name for Anodon rotundum Spix ; 1846, Voy. Amer. Merid., 5, pt. 3, p. 619, listed as a synonym of Anodonta trapezum Spix. BMNH [lost]. staudingeri Ihering, Mycetopus : 1890 Archiv. fur Natur., 56 (i) : 131, figs, a-b (Oberen Ama- zonas, und seiner Zufliisse von Ecuador und Peru). Lectotype Senckenberg Mus. 3802, selected by Haas, 1931, Senckenbergiana 13: 103 ; 3 paratypes BMNH 1902.11.20.3-4 ; 1902,9.2.35 UNIONACEA: TYPES AND FIGURED SPECIMENS 95 stuarti A. Adams and Angus, Unto : 1864, Proc. Zool. Soc. London, p. 417 (lagoon, near Mt. Margaret, Central Australia). Holotype BMNH 1870.10.26.40, figured by McMichael and Hiscock, Australian Jour. Marine and F. W. Res., 9, pi. 2, figs. 10-11. subcrassa Lea, Anodonta : 1850, Proc. Zool. Soc. London, p. 198 ; 1851, Ann. and Mag. Nat. Hist., 8 : 495 ; 1859, Jour. Acad. Nat. Sci. Phila. (2) 4 : 236, pi. 33, fig. 115 ; 1860, Obs. Unio, 7 : 54 (Laguna de Bai, Luzon, Philippine Islands, Cuming colln.). Holotype USNM 86693 .' 2 paratypes BMNH 42.5.10.1529 Cuming colln., one figured by Sowerby, 1877, Conch. Iconica, 17, Anodon, pi. 42, species 42. subjecta Iredale, Centralhyria angasi : 1934, Australian Zool., 8 : 67, based in part on shell figured by Smith, 1874, Zool. of " Erebus " and " Terror " : Mollusca, p. 3, pi. 4, fig. 2. Selected as lectotype BMNH 40.10.21.29, by McMichael and Hiscock, 1958, Australian Jour. Marine and F. W. Res., 9 : 427, pi. 8, figs. 6-7 (Avon River, Western Australia). subreniformis Sowerby, Anodon : 1867, Conch. Iconica, 17, Anodon, pi. 14, species 50 (Lake Nyassa, Cuming colln.). Holotype BMNH 1965192. subsinuatus Sowerby, Anodon : 1867, Conch. Iconica, 17, Anodon, pi. 7, species 14 (Hab. ?, Cuming colln . ) . BMNH [lost] . subtortus Baird and Adams, Unio : 1867, Proc. Zool. Soc. London, p. 491, pi. 26, figs, i, la (Shanghai, North China). Holotype BMNH 1867.5.18.17, refigured by Sowerby 1868, Conch. Iconica, 16, Unio, pi. 87, species 465. subtrigonus Sowerby, Unio : 1867, Conch. Iconica, 16, Unio, pi. 58, species 292 (Siam, Taylor colln.). Holotype BMNH 1874.12.11.3. swainsoni Sowerby, Unio : 1868, Conch. Iconica, 16, Unio, pi. 76, fig. 396 (Hab. ?, Hanley colln.). Holotype BMNH 1900.3.19.21. swinhoei, H. Adams, Anodonta : 1866, Proc. Zool. Soc. London, p. 446 (Formosa). Probable measured holotype BMNH 78.1.28.200 ; smaller paratype BMNH 78.1.28.201. swinhoei Sowerby, Unio : 1866, Conch. Iconica, 16, Unio, pi. 42, species 232 (Camboja, Capt. Swinhoe, Cuming colln.). Holotype BMNH 1968653. syriaca Pallary, Margaritana : 1929, M^m. Inst. d'Egypte, 12, 34, text-fig. (Nahr el-K6bir, 30 km. au Sud de Tartous [Syria]). 2 paratypes BMNH 1937.12.30.13052-3. tabascensis Morelet, Anodonta : 1884, Jour, de Conch., 32 : 124 (Marais du Tabasco [Mexico]). Holotype BMNH 93.2.4.2008, figured by Fischer and Crosse, 1894, Miss. Sci. au Mexique,pt. 7, 2 : 530, pi. 62, fig. i. tabula Sowerby, Anodon : 1867, Conch. Iconica, 17, Anodon, pi. 18, species 68 (Sierra Leone, Cuming colln.). Holotype BMNH 1965190 ; paratype BMNH 1965191. tanganyicensis E. A. Smith, Spatha : 1880, Proc. Zool. Soc. London, p. 350, pi. 31, figs. 8, 8a. (Lake Tanganyika [Central Africa]). Lectotype, here selected, BMNH 80.3.5.48, specimen in figure 8 ; paralectotypes BMNH 80.3.5.49-53. tanganyicensis E. A. Smith, Unio : 1880, Proc. Zool. Soc. London, p. 351, pi. 32, figs. 9, 93. (Lake Tanganyika [Central Africa]). Lectotype, here selected, BMNH 80.3.5.39 specimen in figure 9 ; paralectotypes BMNH 80.3.5.40-47. tanganyicensis E. A. Smith, Unio nyassaensis : 1881, Proc. Zool Soc. London, p. 298, pi. 34, fig. 34a (Lake Tanganyika [Central Africa]). Figured holotype BMNH 1880.12.20.39, consists of one valve : paratypes BMNH 1880.12.20.40-41, consists of two valves ; 3 paratypes BMNH 1880.12.10.130-131, consists of 3 odd valves. tauriformis Fulton, Unio (Cumeopsis) : 1906, Ann and Mag. Nat. Hist. (7) 17 : 246, pi. 9, fig. 9 (Yunnah-fu, Yunnan [China]). Holotype BMNH 1906.5.8.72. tavoyensis Gould, Unio : 1843, Proc. Boston Soc. Nat. Hist., 1 : 140 ([Tavoy] British Burmah [Burma]). Measured holotype MCZ 169389 figured by Johnson, 1964, Bull. United States Natl. Mus., 239, p. 156, pi. 32, fig. 4 ; paratypes BMNH 1965184, largest one figured by Reeve, 1864, Conch. Iconica, 16, Unio, pi. 13, species 49. tennis Lea, Anodonta : 1851, Proc. Zool. Soc. London for 1850, pt. 18, p. 198 (Sual, Luzon, Philippine Islands, Cuming) ; Lea 1851, Ann. Mag. Nat. Hist. (2) 8 : 494 ; Lea, 1860, Jour. Acad. Nat. Sci. Phila. (2) 4 : 232, pi. 33, fig. 116 ; Lea, 1860, Obs. Unio, 7 : 55. Lea had 4 specimens before him. The holotype was not found in the BMNH. Paratype BMNH 96 R. I. JOHNSON 1968654 ; paratype USNM 86764 ; 4 specimens under this name, MCZ 175578, ex Cuming. A specimen was figured by Sowerby, 1867, Conch. Iconica 17, Anodon, pi. 15, species 55, from the Cuming colln. but it was not the holotype. testudineus Morelet, Unio : 1841, Testacea Novissima, 1 : 28 (flumen Usumasinta [Peten Prov., Guatemala]). Measured holotype BMNH 94.3.22.1, figured by Fischer and Crosse, 1894, Miss. Sci. au Mexique, pt. 7, 2 : 571, pi. 62, fig. 3. The specimen figured by Reeve, 1865, Conch. Iconica, 16, Unio, pi. 22, species 101, from the Cuming colln. has Morelet's label, but it is not this species. thotnsoni E. A. Smith, Unio : 1880 Ann. and Mag. Nat. Hist. (5) 6 : 430 (Lake Tanganyika [Central Africa]) ; Smith, 1881, Proc. Zool. Soc. London, p. 299, pi. 34, fig. 36. Holotype BMNH 80.12.20.36 ; 2 paratypes BMNH 80.12.20.37-38. thwait[e]sii Lea, Unio : 1859, Proc. Acad. Nat. Sci. Phila., 11 : 152 (Ceylon, Mr. Thwaites, H. Cuming) ; Lea, 1860, Jour. Acad. Nat. Sci. Phila. (2) 4 : 246, pi. 37, fig. 125 ; Lea, 1860, Obs. Unio, 7 : 64 ; Reeve, 1865, Conch. Iconica, 16, Unio, pi. 23, species 105. Holotype BMNH 1965179. Two smaller specimens not seen by Lea, BMNH 1965180. tortuosus Sowerby, Unio : 1868, Conch. Iconica 16, Unio, pi. 65, species 330 (Maryland, Taylor colln.). Holotype BMNH 74.12.11.25. trapezia Martens, Spat ha : 1897, Bes. Weich. Deutsch-Ost-Afrikas, p. 243, text fig. (Victoria- Nyansa im Siiden bie Bussisi ; im Siidwesten bei Nyemirembe und Nyamagotso ; im Westen bie Bare an der Buddu-Kiiste ; bei Bukoba ; Mhugu an der Ostseite des Sees). Paratype BMNH 1902.5.26.44 from the second locality. trapezialis Orbigny, Iridina : 1835, Mag. de Zool., p. 43 ; 1846, Voyage Ame"r. Merid., 5, pt. 3, P- 596, (les bords du Parana, au-dessus de la ville de Corrientes, et aux environs de la Bajada, province d'Entre-Rios, republique Argentine), non Anodonta trapezialis Lamarck teste Orbigny. Not figured by Orbigny and not found in the BMNH. triangularis Sowerby, Anodon : 1867, Conch. Iconica, 17, Anodon, pi. 15, species 56 : (Hab. ?, Cuming colln.). 1870 Ibid., pi. 29, fig. 5&b. The second specimen figured, was selected as " Lectotype " BMNH 1968211, by Sowerby, and is the only specimen extant. trirostris Benson, Unio : 1863 [in] Hanley, Photographic Conch., pi. 2, fig. 9 (no locality [Hindostan]). Figured holotype BMNH 1907.2.30.45. tristis Morelet, Unio : 1845, Moll, de Portugal, p. 107, pi. 13, fig. 2 (a peu distance d'Amarante, au bord de la Tam6ga [Portugal]). Holotype BMNH 93.2.4.1961. tsadianus Martens, Unio (Grandidieria) 1903, Sitzber. Gesell. Natur. Freunde, Berlin, p. 8 (siidufer des Tsad-Sees [Borneo]). Paratype BMNH 1904.4.29.17. tumens Haas, Anodontites lautus : 1910, Ann. and Mag. Nat. Hist. (8) 6 : 499 (Yamashira, Japan). Holotype Senckenberg Mus. 3671, ex. Sowerby and Fulton ; 2 paratypes BMNH 1912.8. 16.123-4. tutnidus Morelet, Monocondylus : 1866, Jour, de Conch., 14 : 62 (in torrentibus montanis Cambodiae). Holotype BMNH 93.2.4.1734, figured by Morelet, 1875, Series Conch., pt. 4, p. 337, pi. 16, fig. i ; 2 paratypes BMNH 93.2.4.1735 ; 93.2.4.1566. tunizana Morelet, Anodonta : 1864, Jour, de Conch., 12 : 156 (La Calle [Tuniza]). Lectotype, here selected, BMNH 1893.2.4.1964, plate 2 fig. 4. Length 55, height 32, width 20 mm. turcicus " Parreiss " [sic] Kuester, Unio : 1862 [in] Martini and Chemnitz, Conch. Cab. (2) 9, pt. 2, p. 267, pi. 90, figs. 3-4 (Kleinasien). 2 specimens BMNH 47.5.28.1-2, distributed by Parreyss before description by Kuester. unicolor Bernardi, Galatea kochii : 1860, Monog. Galatea et Fischeria, p. 23, pi. 4, figs. 3-4. ([Central Africa] Cuming colln.). BMNH [lost]. vaalensis Chaper, Unio : 1885, Bull. Soc. Zool. France, 10 : 480, pi. n, figs. 1-3 (Le Vaal (Afrique australe) aupres de Barclay a peu distance des mines de diamant du Griqualand West) Paratype BMNH 1937.12.30.1314. vellicatus Reeve, Unio : 1865, Conch. Iconica, 16, Unio, pi. 22, species 103 (Guatemala, Cuming colln.). BMNH [lost]. ventricosus Orbigny, Mycetopus : 1846, Voy. Amer. Me'rid., 5, pt. 3, p. 602, pi. 72, figs. 1-3 UNIONACEA: TYPES AND FIGURED SPECIMENS 97 (province de Chiquitos, republique de Bolivia, aux sources de Rio de Tucabaca, non loin de la mission de San- Juan). BMNH [lost]. verecundus Gould Unio : 1850 Proc. Boston Soc. Nat. Hist., 3 : 295 (Manila [Philippine Islands]) ; Gould 1852, United States. Expl. Exped., 12 : 431, pi. 37, figs. 541, a-c. Figured holotype USNM 5926 ; paratype BMNH 1964150, figured by Reeve 1865, Conch. Iconica, 16, Unio, pi. 25, species 125. verrucifer Martens, Unio : 1894, Sitzber. Gesell. Natur. Freunde, Berlin, p. 216 (Fluss Hangang [bei Soul, Prov. Kyongkwido], Korea) ; Martens, 1905, Zool. Jahrb., Suppl. 8, p. 61, pi. 3, figs. 2, 2b. Paratype BMNH 94.11.24.9, consists of one valve. vignouana Bernardi, Margaritana : 1858, Jour, de Conch., 7 : 302, pi. 10, fig. i (unlacvosinde la riviere Como, Haut Gabon). Figured holotype and two paratypes in colln. of Jour de Conch., Paris teste Fischer, 1950, Jour, de Conch., 90 : 18 ; paratype BMNH [lost] figured by Reeve, 1865, Conch. Iconica, 16, Unio, pi. 25, species 120 as Unio vignonana. vittatus Lea, Unio : 1859, Proc. Acad. Nat. Sci. Phila., 11 : 153 (Australia, H. Cuming) ; Lea, 1860, Jour. Acad. Nat. Sci. Phila. (2) 4 : 249, pi. 38, fig. 128 : Lea, 1860, Obs. Unio, 7 : 67. Opposite valve of type also figured by Reeve, 1864, Conch. Iconica, 16, Unio, pi. 18, species 83, Holotype BMNH 1958.4.3.1 ; paratype USNM 84407. See discussion by McMichael and Hiscock, 1958, Australian Jour, of Marine and F. W. Res., 9 : 390, pi. 2, fig. i, pi. i, figs. 13-14. vonbuschea Sowerby, Unio : 1866, Conch. Iconica, 16, Unio, pi. 51, species 269. (Hab. ?, Cuming colln.). BMNH [lost]. vulcanus Hanley, Unio : 1875, Proc. Zool. Soc. London, p. 606 (Birmah or Pegu). Holotype BMNH 1900.3.19.20, figured by Hanley and Theobald, 1876, Conch. Indica, p. 62, pi. 155, fig. 3- walpolei Hanley, Monocondylaea : 1871, Proc. Zool. Soc. London, p. 587 (Sarawak, Borneo). LectotypeMCZ 175577, selected by Johnson 1948, Nautilus, 62 : 49, pi. 3, fig. 2 ; paralecto- type BMNH 71.7.11.1, ex R. F. Geale. watersoni Tomlin, Anodonta : 1923, Jour, of Conch., 17 : 68, textfig. (Lake Beschik, Salonika). Figured holotype BMNH 1923.12.31.95. welwitschii Morelet, Iridina : 1868, Voyage dans les Royaumes d' Angola et de Benguella, p. 98 (la riviere Muria, pres de Trombeta (Golungo-Alto) Angola). Lectotype, here selected, BMNH 93.2.4.1740, plate i fig. i. Length 86, height 48, width 22 mm ; smaller paralecto- type BMNH 93.2.4.1741. wolwichii Morelet, Unio : 1845, Moll, de Portugal, p. 105, pi. 13, fig. i (les eaux de la val!6e du Tage, entre Villa-Nova et Azumbuja [Portugal]). Holotype BMNH 93.2.4.1964 ; paratype BMNH 93.2.4.1965. woodthorpi Godwin- Austen, Margaritanopsis : 1919, Records Indian Mus., Calcutta, 16 ; 204, pi. 14 (Fort Stedman, Shan States). Holotype BMNH 1965218 ; paratype BMNH 1965219. zatnbesiensis Preston, Unio : 1905, Proc. Mai. Soc. London, 6 : 301, text fig. i (just above Victoria Falls, Zambesi River [Africa]). Figured holotype BMNH 1905.8.29.1. zonata Hanley and Theobald, Unio marginalis : 1872, Conch. Indica, p. 20, pi. 44, fig. 2 (Bel- gaum, Deccan). Holotype BMNH 1907.10.28.252. Part 2 FIGURED SPECIMENS OF UNIONACEA IN THE BRITISH MUSEUM (NAT. HIST.) WHICH ARE NOT TYPES Material in square brackets indicates additional information, or more generally, that the locality was copied from the original description and does not necessarily apply to the specimen figured. Specimens of Unionacea, exclusive of types, figured in : Reeve, L. and G. B. Sower- by, 1864-70, Conchologia Iconica, 16, 17. 1864-68, 16, Unio. 98 R. I. JOHNSON PI. I, fig. I. Unio boykinianus Lea. [Chattachoochee River, Georgia and Warrier River, Alabama]. Cuming colln. BMNH 196473. PI. i, fig. 3. Unio schooler aftensis Lea. [Fox River, Green Bay, Wisconsin, N. America]. Cuming colln. BMNH 196474. PL I, fig. 4. Unio verrucosus Rafinesque. [River Ohio, N.America]. Cuming colln. BMNH 196475. PL 2, fig. 6. Unio kleinianus Lea. Suwanee River, Florida, North America. Cuming colln. BMNH 196476. PL 2, fig. 7. Unio caelatus Conrad. North America. Cuming colln. BMNH 196477. PL 3, fig. 10. Unio turgidus Lea. New Orleans, North America. Cuming colln. BMNH 196478. PL 3, fig. ii. Unio apiculatus Say. North America. Cuming colln. BMNH 196479. PL 4, fig. 14. Unio sparsus Lea. East Tennessee, North America. Cuming colln. BMNH 196480. PL 4, fig. 15. Unio phillipsii Conrad. North America. Cuming colln. BMNH 196481. PL 4, fig. 16. Unio costatus Rafinesque. North America. Cuming colln. BMNH 196482. PL 5, fig. 17. Unio trapezoides Lea. [Lake St. Joseph, Louisiana, North America]. Cuming colln. BMNH 196483. PL 5, fig. 20. Unio elliotii Lea. [Othcalooga Creek, Gordon County, Georgia, North America.] Cuming colln. BMNH 196484. PL 6, fig. 22. Unio flexuosus Rafinesque. North America. Cuming colln. BMNH 196485. PL 6, fig. 24. Unio quadratus Rafinesque. Ohio River, North America. Cuming colln. BMNH 196486. PL 7, fig. 25. Unio metanever Rafinesque. North America. Cuming colln. BMNH 196487. PL 7, fig. 26. Unio prasinus Conrad. North America. Cuming colln. BMNH 196488 and 196489. PL 7, fig. 27. Unio fragosus Conrad. North America. Cuming colln. BMNH 196490. PL 8, fig. 28. Unio cyphius Rafinesque. North America. Cuming colln. BMNH 196491. PL 8, fig. 29. Unio dromas Lea. Tennessee, North America. Cuming colln. BMNH 196492. PL 9, fig. 32. Unio nodulosus Wood. [Unio leeai Gray (Errata)]. China. Cuming colln. BMNH 196493. PL 9, fig. 34. Unio graniferus Lea. Ohio River, North America. Cuming colln. BMNH 196494. PL 9, fig. 35. Unio perplicatus Conrad. North America. Cuming colln. BMNH 196495. PL 10, fig. 36. Unio semigranosus Philippi. Mexico. Cuming colln. BMNH 196496. PL 10, fig. 38. Unio dorfeuillianus Lea. Ohio River, North America. Cuming colln. BMNH 196497. PL ii, fig. 40. Unio hippopaeus Lea. Lake Erie, North America. Cuming colln. BMNH 196498. PL ii, figs. 4ia, b. Unio gibbosus Rafinesque. North America. Cuming colln. a is BMNH 196499 ; b is BMNH 1964104. PL ii, fig. 42. Unio perlensis Conrad. North America. Cuming colln. BMNH 1964100. PL 12, fig. 43. Unio bullatus Rafinesque. North America. Cuming colln. BMNH 1964101. PL 12, fig. 45. Unio stegarius Rafinesque. North America. Cuming colln. BMNH 1964102. PL 12, fig. 46. Unio pernodosus Lea. North Carolina, North America. Cuming colln. BMNH 1964103. PL 13, fig. 47. Unio nodiferus Conrad. [North America]. Cuming colln. BMNH 1964105. PL 13, fig. 48. Unio intermedius Conrad. North America. Cuming colln. BMNH 1964106. PL 13, fig. 52. Unio stapes Lea. Alabama, North America. Cuming colln. BMNH 1964107. PL 14, fig. 54. Unio rangianus Lea. Ohio River, North America. Cuming colln. BMNH 1964108. PL 14, fig. 55. Unio penitus Conrad. Alabama, North America. Cuming colln. BMNH 1964109. UNIONACEA: TYPES AND FIGURED SPECIMENS 99 PL 14, fig. 57. Unio arcaeformis Lea. Tennessee River, North America. Cuming colln. BMNH 1964110. PL 14, fig. 58. Unio sowerbyanus Lea. Tennessee [River], North America. Cuming colln. BMNH 1964111. PL 14, fig. 59. Unio pileus Lea. Ohio River, North America. Cuming colln. BMNH 1964112. PL 14, fig. 60. Unio sulcatus Lea. Ohio River, North America. Cuming colln. BMNH 1964113. PL 15, fig. 62. Unio haysianus Lea. Cumberland River, North America. Cuming colln. BMNH 1964114. PL 15, fig. 63. Unio ridibundus Say. North America. Cuming colln. BMNH 1964115. PL 15, fig. 64. Unio personatus Say. Ohio River, North America. Cuming colln. BMNH 1964116. PL 15, fig. 65. Unio edgarianus Lea. Tennessee River, North America. Cuming colln. BMNH 1964117. PL 15, fig. 66. Unio stewardsonii Lea. Chattanooga River, North America. Cuming colln. BMNH 1964118. PL 16, fig. 68. Unio nucleopsis Conrad. North America. Cuming colln. BMNH 1964119. PL 16, fig. 69. Unio chattanoogaensis Lea. Chattanooga, [River] North America. Cuming colln. BMNH 1964120. PL 16, fig. 70. Unio raveneliensis Lea. Tennessee River, North America. Cuming colln. BMNH 1964121. PL 16, fig. 71. Unio decisus Lea. Alabama River, North America. Cuming colln. BMNH 1964122. PL 16, fig. 72. Unio mundus Lea. North America. Cuming colln. BNMH 1964123. [Not the mundus of Lea, but nearer his mooresianus and cuneolus (Errata)]. PL 16, fig. 73. Unio cuneatus Rafinesque. North America. Cuming colln. BMNH 1964124. PL 1 6, fig. 74. Unio woodwardianus Lea. Georgia, North America. Cuming colln. BMNH 1964125. PL 17, fig. 75. Uniolenior'Lea.. Tennessee, North America. Cuming colln. BMNH 1964126. PL 17, fig. 77. Unio medius Lea. Alabama, North America. Cuming colln. BMNH 1964127. PL 17, fig. 78. Unio subrostratus Say. North America. Cuming colln. BMNH 1964128. [Nearer the patulus of Lea (Errata)]. PL 17, fig. 79 a. Unio capsaefromis Lea. Cumberland River, North America. Cuming colln. BMNH 1964129. PL 18, fig. 92. Unio perdix Lea. Tennessee, North America. Cuming colln. BMNH 1964130. PL 18, fig. 84. Unio obesus Lea. [York River, Virginia] North America. Cuming colln. BMNH 1964131. PL 19, fig. 85. Unio mississippiensis Conrad, MS. Mississippi River, North America. Cuming colln. BMNH 1964132. Sent by Anthony to Cuming prior to publication in 1850. PL 19, fig. 86. Unio rectus Lamarck. North America. Cuming colln. BMNH 1964133. PL 19, fig. 87. Unio anodontoides Lea. North America. Cuming colln. BMNH 1964134. PL 20, fig. 88. Unio buddianus Lea. Florida, North America. Cuming colln. BMNH 1964135. PL 20, fig. 89. Unio cucumoides Lea. Hunter's River, New South Wales. Cuming colln. BMNH 1964136. PL 20, fig. 90. Unio shepardianus Lea. Georgia, North America. Cuming colln. BMNH 1964137. PL 21, fig. 93. Unio patagonicus d'Orbigny. Patagonia. Cuming colln. BMNH 1964138. PL 21, fig. 94. Unio nasutus Say. North America. Cuming colln. BMNH 1964139. PL 21, fig. 95. Uniofuscatus Lea. Florida, North America. Cuming colln. BMNH 1964140. PL 23, fig. 107. Unio cuneolus Lea. Tennessee, North America. Cuming colln. BMNH 1964142. PL 23, fig. 108. Unio berlandierii Lea. Mexico. Texas. Cuming colln. BMNH 1964143. PL 23, fig. 109. Unio altilis Conrad. North America. Cuming colln. BMNH 1964144. ioo R. I. JOHNSON PL 24, fig. 113. Unio fisherianus Lea. [Chester River, Maryland, North America]. Cuming colln. BMNH 1964145. PL 24, fig. 114. Unio navigioliformis Lea. Hab.? Cuming colln. BMNH 1964146. PL 24, fig. 115. Unio purpuratus Lamarck. Louisiana, North America. Cuming colln. BMNH 1964147. PL 24, fig. 116. Unio haleianus Lea. [Mississippi, near New Orleans, North America]. Cum- ing colln. BMNH 1964148. PL 25, fig. 119. Unio grayanus Lea. China. Cuming colln. BMNH 1964149. PL 25, fig. 125. Unio verecundus Gould. Manila, Island of Luzon, Philippines. Cuming colln. BMNH 1964150. PL 26, fig. 126. Unio ingallsianus Lea. [Siam ; Dr. Ingalls]. Cuming colln. BMNH 1964151. PL 26, fig. 127. Unio incrassatus Lea. Georgia, North America. Cuming colln. BMNH 1964152. PL 26, fig. 128. Unio concestator Lea. Georgia, North America. Cuming colln. BMNH 1964153. PL 26, fig. 129. Unio obtusus Lea. Georgia, North America. Cuming colln. BMNH 1964154. PL 26, fig. 132. Unio aegyptiacus Cailliaud. [River Nile, Egypt ; Cailliaud]. Cuming colln. BMNH 1964155. PL 27, fig. 133. Unio solidus Lea. [Ohio River, at Cincinnati, Ohio, North America]. Cuming colln. BMNH 1964156. PL 27, fig. 135. Unio circulus Lea. Ohio and Tennessee, North America. Cuming colln. BMNH 1964157. PL 27, fig. 139. Unio radiatus Gmelin. [Lake Georgia, North America]. Cuming colln. BMNH 1964158. PL 28, fig. 142. Unio sumatrensis Lea. Sumatra. Cuming colln. BMNH 1964159. This specimen was probably not seen by Lea. Holotype USNM 84059. PL 28, fig. 144. Unionitens~Lza,. Tennessee, North America. Cuming colln. BMNH 1964160. PL 29, fig. 148. Unio iris Lea. Ohio. Cuming colln. BMNH 1964162. PL 29, fig. 149. Unio contradens Lea. River Mezherdeh, Tunis. Cuming colln. BMNH 1964163. PL 30, fig. 154. Unio platyrhynchus Rossmassler. Carinthia. Cuming colln. BMNH 1964164. PL 30, fig. 156. Unio aucklandicus Gray, Auckland, New Zealand, Cuming colln. BMNH 1964165. PL 31, fig. 162. Unio ornatus Conrad. Alabama. [Cuming colln.]. BMNH 1964166. PL 31, fig. 163. Unio subovatus Lea. Ohio. [Cuming colln.]. BMNH 1964167. PL 31, fig. 164. Unio ovatus Say. Ohio. [Cuming colln.]. BMNH 1964168. PL 32, fig. 165. Unio negatus Lea. [Big Prairie Creek, Alabama ; E. R. Showalter. Columbus ; Miss. ; Spillman]. Cuming colln. BMNH 1964169. PL 32, fig. 167. Unio shuttleworthi Lea. Australia ; Shuttleworth. Cuming colln. BMNH 1964170. Not seen by Lea, but mentioned by him as being in the Cuming colln. Holotype USNM 84416. PL 32, fig. 169. Unio burroughianus Lea. River Panama, South America. Cuming colln. BMNH 1964171. PL 33, fig. 170. Unio umbrosus Lea. [Medellin River, Mexico ; Dr. Burrough]. Cuming colln. BMNH 1964172. PL 33, fig. 171. Unio medellinus Lea. [River Medellin, near Vera Cruz ; Dr. Burrough. Cuming colln.]. BMNH 1964173. PL 33, fig. 172. Unio hembeli Conrad. Louisiana. Cuming colln. BMNH 1964174. PL 33, fig. 173. Unio lecontianus Lea. [River Georgia ; Major Leconte]. Cuming colln. BMNH 1964175. PL 34, fig. 175. Unio buckleyi Lea. [Lake George and Lake Munroe Florida: S.B.Buckley]. Cuming colln. BMNH 1964176. UNIONACEA: TYPES AND FIGURED SPECIMENS 101 PL 34, fig. 176. Unio discrepans Lea. [North Alabama ; Prof. Tuomey]. Cuming colln. BMNH 1964177. PI. 34, fig. 179. Unio umbrans Lea. [Othcalooga Creek, Gordon County, Georgia ; Bishop Elliot]. Cuming colln. BMNH 1964178. PI. 34, fig. 180. Unio barnesianus Lea. [Cumberland River, Tennessee ; Dr. Troost]. Cuming colln. BMNH 1964179. PL 34, fig. 181. Unio regularis Lea. [French Broad River, East Tennessee ; Dr. Troost]. Cuming colln. BMNH 1964180. PL 35, fig. 182. Unio micans Lea. [Catawba River ; Wheatley. Deep River Gulf, North Carolina, Emmons. Cuming colln.]. BMNH 1964181. PL 35, fig. 183. Unio pellucidus Lea. [Chatahoochee River, Georgia]. Cuming colln. BMNH 1964182. PL 35, fig. 184. Unio heterodon Lea. Georgia. Cuming colln. BMNH 1964183. PL 35, fig. 185. Unio confertus Lea. South Carolina. Cuming colln. BMNH 1964184. PL 35, fig. 186. Unio parvus Barnes. [Fox River, North America]. Cuming colln. BMNH 1964185. PL 35, fig. 187. Unio blandingianus Lea. Florida. Cuming colln. BMNH 1964186. PL 35, fig. 1 88. Unio haslehurstianus Lea. [Satilla River, Camden County, Georgia ; C. T. Downie]. Cuming colln. BMNH 1964187. PL 36, fig. 189. Unio acutissimus Lea. Alabama River. Cuming colln. BMNH 1964188. PL 36, fig. 190. Unio glans Lea. River Ohio. [Cuming colln.]. BMNH 1964189. PL 36, fig. 191. Unio zeiglerianus Lea. Cumberland River, Tennessee. Cuming colln. BMNH 1964190. PL 36, fig. 192. Unio hildrethianus Lea. Ohio. Cuming colln. BMNH 1964191. PL 36, fig. 193. Unio tortivus Lea. [Chattahoochee River, Georgia]. Cuming colln. BMNH 1964192. PL 36, fig. 194. Unio ahaeneus Lea. [Black Creek, Florida]. Cuming colln. BMNH 1964193. PL 36, fig. 196. Unio fabalis Lea. [Ohio River ; Lea]. Cuming colln. BMNH 1964194. PL 36, fig. 197. Unio cumberlandicus Lea. Cumberland River. Cuming colln. BMNH 1964195. PL 37, fig. 200. Unio lens Lea. [River Ohio ; Lea. Tennessee ; Vanuxem]. Cuming colln. BMNH 1964196. PL 37, fig. 201. Unio subrotundus Lea. West Africa. [Cuming colln.]. BMNH 1964197. PL 37, fig. 202. Unio foremanianus Lea. Coosa River, Alabama. Cuming colln. BMNH 1964198. PL 37, fig. 203. Unio hydeanis Lea. [Teche River, Louisiana ; W. M. Stewart]. Cuming colln. BMNH 1964199. PL 37, fig. 204. Unio argenteus Lea. [Holston River, East Tennessee ; Dr. Troost]. Cuming colln. BMNH 1964200. PL 38, fig. 206. Unio novi-eboraci Lea. [Oak Orchard Creek, Orleans County, New York ; J.C. Jay]. Cuming colln. BMNH 1964201. PL 38, fig. 207. Unio murchisonianus Lea. [Unio douglasiae Gray (Errata)]. China. Cuming colln. BMNH 1964202. PL 38, fig. 208. Unio exiguus Lea. Chatahoochee River, Georgia. Cuming colln. BMNH 1964203. [Not so, possibly fatuus (Errata)]. PL 38, fig. 210. Unio troostensis Lea. North America. Cuming colln. BMNH 1964204. PL 38, fig. 211. Unio muhlfeldianus Lea. Tennessee. Cuming colln. BMNH 1964205. PL 38, fig. 212. Unio obesus Lea. [York River]. Cuming colln. BMNH 1964206. PL 39, fig. 213. Unio ziczac Lea. South Carolina. Cuming colln. BMNH 1964207. PL 39, fig. 215. Unio gracilis Barnes. River Ohio. Cuming colln. BMNH 1964208. PL 39, fig. 216. Unio vanuxemensis Lea. River Cumberland, Tennessee. Cuming colln. BMNH 1964209. PL 39, fig. 217. Unio stramineus Conrad. River Alabama. Cuming colln BMNH 1964210. 102 R. I. JOHNSON PI. 40, fig. 219. Unio gibber Lea. [Carryfork River, Tennessee ; Prof. Troost]. Cuming colln. BMNH 1964211. PI. 40, fig. 220. Unio crassus Say. North America. Cuming colln. BMNH 1964212. PI. 40, fig. 221. Unio infucatus Conrad. Georgia, America. Cuming colln. BMNH 1964213. PL 41, fig. 223. Unio fibuloides Lea. Connasuaga River, Georgia. Cuming colln. BMNH 1964214. PL 41, fig. 226. Unio coffer Krauss [Is U. africanus Lea (Errata)]. Natal. Cuming colln. BMNH 1964215. PL 41, fig. 227. Unio bigbyensis Lea. Big Bigby Creek, Tennessee. Cuming colln. BMNH 1964216. PL 41, fig. 228. Unio dolosus Lea. Alabama River. Cuming colln. BMNH 1964217. PL 42, fig. 233b. Unio lindsleyi Lea. [Tennessee ; President Lindsley]. Cuming colln. BMNH 1964218. PL 43, fig. 237. Unio delphinulus Morelet. Yucatan. Hanley colln. BMNH 1917.10.28.182. PL 44, fig. 241. Unio schwartzenbachii Rossmassler [Bourguignat]. Hab.? Hanley colln. BMNH 1907.10.28.197. PL 47, fig. 253. Unio greenii Conrad. Virginia. Hanley colln. BMNH 1907.10.28.192. PL 47, fig. 254. Unio hainesianus Lea. Little Arkansas. Cuming colln. BMNH 1964219. PL 47, fig. 255. Unio postellii Lea. [Randall's Creek, near Columbia, Georgia]. Cuming colln. BMNH 1964220. PL 48, fig. 257. Unio leptodon Rafinesque. Hab. ?. Cumin colln. BMNH 1964354. PL 48, fig. 259. Unio arcula Lea. Georgia, in America. Cuming colln. BMNH 1964355. PL 48, fig. 260. Unio housei Lea. Georgia, in America. [Siam]. Cuming colln. BMNH 1964356- PL 49, fig. 261. Unio spinosus Lea. Georgia, in America. Cuming colln. BMNH 1964357. PL 49, fig. 262. Unio crispisulcatus Lea. [Benson (Errata)]. Hab? Cuming colln. BMNH 1964358. PL 49, fig. 263. Unio collinus Lea [Conrad]. Virginia. Cuming colln. BMNH 1964359. PL 50, fig. 265. Unio myersianus Lea. Camboja, Lao Mountains [Siam]. Cuming colln. BMNH 1964361. PL 50, fig. 266. Unio complanatus Barnes. Ohio. Cuming colln. BMNH 1964362. PL 51, fig. 267. Unio marginatus Lea. [Say] Massachusetts. Cuming colln. BMNH 1964363. PL 51, fig. 270. Unio poulsoni Conrad. Mississippi. Cuming colln. BMNH 1964364. PL 52, fig. 271. Unio gravidus Lea. Siam. Cuming colln. BMNH 1964365. PL 53, fig. 275. Unio discoideus Lea. Hab. ?. Cuming colln. BMNH 1964366. PL 53, fig. 276. Unio nicklinianus Lea. China. Cuming colln. BMNH 1964367. PL 54, fig. 277. Unio percoarctatus Lea. [Not so, approaches laevissimus Lea (Errata)]. Hab.? Cuming colln. BMNH 1964368. PL 56, fig. 284. Unio decurvatus Rossmassler. Germany. Taylor colln. BMNH 1874.12.11. 14. PL 57, fig. 290. Unio funebralis Lea. Uruguay, South America. Taylor colln. BMNH 1874.12.11.10. PL 57, fig. 291. Unio tampicoensis Lea. Tampico and Tecomate, in Mexico. Taylor colln. BMNH 1874.12.11 2 PL 58, fig. 293 a, b. Unio luteolus Lamarck. Hab. ? Taylor colln. a is BMNH 1874.12.11.18 ; b is BMNH 1874.12.11.19. PL 58, fig. 294. Unio cariosus Say. Hab.?. Taylor colln. BMNH 1874.12.11.7. PL 59, fig. 296. Unio congaraeus Lea. Congaree River, Columbia [South Carolina]. Taylor colln. BMNH 1874.12.11.24. PL 59, fig. 297. Unio marginalis Lamarck. Bengal. Taylor colln. BMNH 1874.12.11.9. PL 59, fig. 298. Unio dolabraeformis Lea. Georgia. Taylor colln. BMNH 1874.12.11.6. PL 60, fig. 299. Unio confragosus Say. North America. Taylor colln. BMNH 1874.12.11.26. PL 60, fig. 301. Unio jayensis Lea. Hab.? Taylor colln. BMNH 1874.12.11.12. PL 60, fig. 302. Unio rugosus Barnes. North America. Taylor colln. BMNH 1874.12.11.20. UNIONACEA: TYPES AND FIGURED SPECIMENS 103 PI. 60, fig. 306. Unio multiradiatus Lea. [Unio luteolus var. (Errata)]. River Ohio. Taylor colln. BMNH 1874.12.11.28. PI. 61, fig. 309. Unio lineatus Lea. [Chattahoochee River, Columbus, Georgia ; Dr. Boykin]. Taylor colln. BMNH 1874.12.11.27. PI. 62, fig. 310. Unio discus Lea. India. [Habitat wrong (Errata)] . So werby colln. BMNH 1888.12.4.2016. PI. 62, fig. 311. Unio sinuatus Lamarck. Rivers of Europe. Taylor colln. BMNH 1864.12. 11.29. PI. 63, fig. 317. Unio ochraceus Say. [Rivers Georgia, Schuylkill, Delaware, Savannah]. Taylor colln. BMNH 1874.12.11.5. PI. 66, fig. 335. Unio pullatus Lea. River Georgia. Taylor colln. BMNH 1874.12.11.16. PL 67, fig. 339. Unio wynegungensis Lea. [Wynegunga River, Nagpoor, Bengal]. Taylor colln. BMNH 1874.12.11.8. PI. 67, fig. 341. Unio roanoakensis Lea. Virginia. Taylor colln. 1874.12.11.21. PL 68, fig. 346. Unio purpureus Say. Massachusetts. Taylor colln. BMNH 1952.10.30.70. PL 68, fig. 349. Unio hopetownensis Lea. [Hopetown, Georgia]. Taylor colln. BMNH 1874.12.11.23. PL 68, fig. 350. Unio downei [sic] Lea. Hopetown, Georgia. Taylor colln. BMNH 1874.12. 11.22. PL 69, fig. 353. Unio osbeckii Philippi. China. Taylor colln. BMNH 1874.12.11.13. PL 69, fig. 354. Unio clava Lamarck. Western waters in N. America. Hanley colln. BMNH 1952.10.30.69. PL 70, fig. 356. Unio camptodon Say. New Orleans, N. America. Hanley colln. BMNH 1952.10.30.68. PL 70, fig. 357. Unio clairbornensis Lea. [Alabama River, near Clairborne, N. America]. Hanley colln. BMNH 1952.10.30.71. PL 70, fig. 358. Unio platyrrhinchoideus Dupuy. Arcachon, Western France. Hanley colln. BMNH 1907.12.30.368. PL 70, fig. 359. Unio powellii Lea. [Saline River, Arkansas, N. America]. Hanley colln. BMNH 1952.10.30.57. PL 71, fig. 360. Unio corrugatus Chemnitz [Miiller]. Coromandel, etc., East Indies. Hanley Colln. BMNH 1907.12.30.65. PL 71, fig. 361. Unio forbesianus Lea. Savannah River, North America. Hanley colln. BMNH 1907.10.28.200. PL 72, fig. 366. Unio ravistellus Morelet. Lake Ysabel, Guatemala. Hanley colln. BMNH 1907.12.30.379. PL 72, fig. 367. Unio striatulus Lea. Roanoke River, N. Carolina. U. States, N. America. Hanley colln. BMNH 1907.10.28.251. PL 72, fig. 369. Unio rotundus Spix. Brazil. Hanley colln. BMNH 1907.10.28.180. PL 72, fig. 370. Unio subtentus Say. S. Carolina, N. America. Hanley colln. BMNH 1907.12.30.313. PL 72, fig. 371. Unio reniformis Schmidt. [Rossmassler]. S. Germany. Hanley colln. BMNH 1907.12.20.371. PL 73, fig. 373. Unio nux persica Dunker. China. Hanley colln. BMNH 1907.10.28.174. PL 73, fig. 374. Unio niloticus Caillaud. The river Nile. Hanley colln. BMNH 1907.10.28. 247. PL 73, fig. 378. Unio phaseolus Hildreth. River Wabash, etc., North America. Hanley colln. BMNH 1952.10.30.132. PL 73, fig. 379. Unio gibbosus Barnes. Upper Mississippi and Missouri, North America. Hanley colln. BMNH 1964369. PL 74, fig. 381. Unio variabilis Maton. Rio de la Plata, S. America. Hanley colln. BMNH 1907.10.28.199. PL 74, fig. 382. Unio ellipticus Spix. Bahia, Brazil. Hanley colln. BMNH 1907.10. 28.198. 104 R- I. JOHNSON PL 74, fig. 383. Unio oregonensis Lea. Columbia River, Oregon. Hanley colln. BMNH 1907.12.30.52. PL 74, fig. 385. Unio calamitarum Morelet. Baluntie, near Palenque. Hanley colln. BMNH 1907.12.30.377. PL 75, fig. 387. Unio pliciferus Lea. Mexico. Hanley colln. BMNH 1097.12.30.375. PL 75, fig. 389. Unio teretiusculus Philippi. Sennaar. Hanley colln. BMNH 1907.10. 28.172. PL 75, fig. 391. Uniofamelicus Gould. Walla Walla, Oregon, North America. Hanley colln. BMNH 1907.12.30.369. PL 75, fig. 392. Unio capigliolo Payraudeau. Corsica, etc. Hanley colln. BMNH 1952. 10.30.133. PL 75, fig. 393. Unio dehiscens Say. Ohio. Hanley colln. BMNH 1907.12.30.370. PL 76, fig. 394. Unio fabula Lea. Cumberland River, Tennessee. Hanley colln. BMNH 1907.10.28.245. PL 76, fig. 397. U nio elongatus Lamarck. Europe. Hanley colln. BMNH 1907.12.30.345. PL 76, fig. 398. Unio holstonianus Lea. Holston River, U.S. North America. Hanley colln. BMNH 1907.12.30.372. PL 76, fig. 399. Unio undulatus Say. Massachusetts, etc., North America. Hanley colln. BMNH 1907.12.30.378. PL 77, fig. 401. Unio corrianus Lea. Pegu, (W. Theobald). Hanley colln. BMNH 1907.10. 28.165. PL 77, fig. 402. Unio kirtlandianus Lea. Ohio, N. America. Hanley colln. BMNH 1907.10. 28.177. PL 77, fig. 403. Unio excavatus Lea. Georgia and Alabama, N. America. Hanley colln. BMNH 1907.12.30.137. PL 78, fig. 406. Unio troostensis Lea. Cumberland River, N. America. Hanley colln. BMNH 1907.10.28.190. PL 78, fig. 407. Unio striatus Lea. Chattahoochie River, Columbus, Georgia, N. America. Hanley colln. BMNH 1907.12.30.57. PL 78, fig. 410. Unio calceola Lea. Ohio. Hanley colln. BMNH 1907.01.28.253. PL 78, fig. 411. Unio corvus Lea. Georgia, N. America. Hanley colln. BMNH 1907.10. 28.186. PL 79, fig. 413 a. Unio sandrii Villa [Rossmassler]. Dalmatia. Hanley colln. BMNH 1952.10.30.135. PL 79, fig. 414. Unio bonellii Ferussac [Rossmassler]. Illyria. Hanley colln. BMNH 1907.10.28.179. PL 80, fig. 418. Uniofallax Lea. Georgia and Tennessee, N. America. Hanley colln. BMNH 1907.10.28.191. PL 80, fig. 419. Unio triangulata Lea. Georgia, N. America. Hanley colln. BMNH 1907. 10.28.246. PL 80, fig. 420. Unio japanensis Lea. Japan. Hanley colln. BMNH 1907.12.30.373. PL 80, fig. 423. Unio lugubris Lea. Hopeton, near Darien, N. America. Hanley colln. BMNH 1907.12.30.39. PL 85, fig. 450. Unio asperatus Lea. Alabama river, N. America. Hanley colln. BMNH 1907.10.28.160. PL 85, fig. 452. Unio glaber Lea. Tennessee, N. America. Hanley colln. BMNH 1907.12. 30.55. PL 85, fig. 454. Unio arctior Lea. Ohio, near Cincinnati, N. America. Hanley colln. BMNH 1907.12.30.59. PL 85, fig. 455. Unio multistriatus Lea. Brazil. Hanley colln. BMNH 1907.10.28.196. PL 85, fig. 456. Unio subovatus Lea. Ohio. Hanley colln. BMNH 1907.10.28.167. PL 86, fig. 457. Unio pellis-lacerti Morelet. Siam. Hanley colln. BMNH 1907.10.29.248. PL 86, fig. 462. Unio stonensis Lea. Stones's River, Tennessee, America. Hanley colln. BMNH 1907.10.28.188. UNIONACEA: TYPES AND FIGURED SPECIMENS 105 PL 86, fig. 463. Unio hochstetteri Dunker. [Lake Taupo, N. Zealand (teste Dunker)]. Hanley colln. BMNH 1907. 10.28.265. PL 87, fig. 466. Unio fontaineanus Orbigny. S. America. Hanley colln. BMNH 1907.10. 28.163. PL 94, fig. 512. Unio coccineus Hildreth [Lea]. Ohio, N. America. BMNH 1907.10.28.238. PL 94, fig. 513. Unio salwenianus Gould. River Salwen British Birmah. Hanley colln. BMNH 1907.10.28.189. PL 95, fig. 516. Unio pugio Benson. E.Indies. Hanley colln. BMNH 1907.10.28.257. PL 95, fig. 517. Unio crebristriatus Anthony. British Birmah, (Theobald). Hanley colln. BMNH 1907.12.30.42. PL 95, fig. 519. Unio peguensis Anthony. Pegu (Theobald). Hanley colln. BMNH 1907. 10.28.259. PL 95, fig. 520. Unio crassus Say. N. America. Hanley colln. BMNH 1907.10.28.262. 1868, 16, Mycetopus. PL i, fig. i. Mycetopus soleniformis Orbigny. Bolivia. Cuming colln. BMNH 1963403. PL 2, fig. 6. Mycetopus emarginatus Lea. Siam. Cuming colln. BMNH 1964404. 1867-70, 17, Anodon. PL 5, fig. ii. Anodon suborbiculatus Say. River Oregon. Cuming colln. BMNH 1964371. PL 6, fig. 13. Anodon gibba [gibbuni] Benson. River Kiang. Cuming colln. BMNH 1964372. PL 8, fig. 15, species 16. Anodon sinuosus Lamarck. Brazil. Cuming colln. BMNH 1964373. PL 8, fig. 1 8, species 19. Anodongigantea'Lea. Port Gibson. Cuming colln. BMNH 1964374. PL 9, fig. 21, species 22. Anodon susannae Griffiths. Rio de la Plata. Cuming colln. BMNH I964375- PL 9, fig. 22, species 23. Anodon wahlamatensis Lea. Rio Sacramento, Wahlamat. Cuming colln. BMNH 1964377. PL 9, fig. 24. Anodon gibbosus Say. River Georgia. Cuming colln. BMNH 1964378. PL 12, fig. 36. Anodon politus [polita~\ Mousson. Siam. Cuming colln. BMNH 1964380. PL 12, fig. 37. Anodon charpentieri Kiister. Hab.? Cuming colln. BMNH 1964381. PL 12, fig. 39. Anodon obtusus Spix. Brazil. Cuming colln. BMNH 1964382. PL 14, fig. 51. Anodon ferussaciana Lea. [Sciato [Scioto] River, near Columbus, Ohio]. Cuming colln. BMNH 1964383. PL 15, fig. 54. A nodon oblongus de Millet. D'Angere, Maine et Loire. Cuming colln. BMNH 1964385. PL 17, fig. 59. Anodon incertus Lea. River Ohio. Cuming colln. BMNH 1964386. PL 17, fig. 60. Anodon edentulus Lea. [Say]. North America. Cuming colln. BMNH 1964387. PL 17, fig. 61. A nodon fragilis Lamarck. River Ohio. Cuming colln. BMNH 1964388. PL 17, fig. 62. Anodon newtonensis Lea [Newtown Creek, New Jersey, near Philadelphia, Pennsylvania] Cuming colln. BMNH 1964389. Identified by Lea, but not a primary type. PL 17, fig. 64. Anodon lucasii Morelet [Deshayes]. Algeria. Cuming colln. BMNH 1964390. PL 1 8, fig. 69. [By Error]. Anodon ferussaciana Lea. Ponds near Montreal, [Canada]. Cuming colln. BMNH 1964391. PL 19, fig. 72. Anodon rayi Dupuy. Hab.? Cuming colln. BMNH 1964393. PL 19, fig. 73. Anodon lacustris Lea. Lakes in New York. Cuming colln. BMNH 1964394. PL 19, fig. 74. Anodon salmonea Lea. Ohio. Cuming colln. BMNH 1964395. PL 27, fig. 102. Anodon imbecillis Lea. Ohio River. Walpole colln. BMNH 1964396. PL 32, fig. 128. Anodon pavonia Lea. [Little Beaver, Ohio]. BMNH 1964399. PL 33, fig. 133. Anodon stewartianus Lea. [River Teche, Louisiana]. BMNH 1841.4.6.24. 1869, 17, Hyria. PL 3, fig. 6. Hyria avicularis Lamarck. Hab.? BMNH 1846.3.6.31. 106 R. I. JOHNSON Specimens of Unionacea, exclusive of types, figured in : Hanley, S., 1842-56, An Illustrated and Descriptive Catalogue of Recent Bivalve Shells, forming an appendix to the Index Testaceo- logicus. Supplementary plates : PI. 20, fig. 20. Unio perdix Lea. Tennessee. BMNH 1907.10.28.267. PL 20, fig. 21. Unio hopetonensis Lea. Near Darien, [Georgia]. BMNH 1907.12.30.54. PI. 20, fig. 22. Unio cariosus Say. N. America. BMNH 1952.10.30.62. PI. 20, fig. 23. Unio circulus Lea. Ohio, etc. BMNH 1907.10.28.243. PI. 20, fig. 26. Unio undulatus Barnes. Ohio. BMNH 1907.12.30.38. PI. 20, fig. 27. Unio tuberculatus Barnes. N. America. BMNH 1907.10.28.162. PL 20, fig. 29. Unio caelatus Conrad. Tennessee. BMNH 1907.12.30.50. PL 20, fig. 32. Unio pliciferus Lea. Mexico. BMNH 1907.10.28.181. PL 20, fig. 33. Unio semigranosus Von dem Busch. Mexico. BMNH 1907.10.28.261. PL 20, fig. 36. Unio tampicoensis Lea. Mexico. BMNH 1907.10.28.268. PL 20, fig. 42. Unio tenuissimus Lea. Ohio. BMNH 1952.10.30.136. PL 20, fig. 44. Unio delphinus Gruner. Mallacca. BMNH 1952.10.30.61. PL 20, fig. 48. Unio ochraceus Say. Delaware. BMNH 1907.10.28.166. PL 20, fig. 51. Unio securis Lea. Ohio. BMNH 1907.12.30.51. PL 20, fig. 53. Unio marginalis Lamarck. Bengal. BMNH 1952.10.30.65. PL 20, fig. 54. Unio gibbosus Barnes. N. America. BMNH 1907.12.30.34. PL 20, fig. 55. Unio splendidus Lea. Georgia in America. BMNH 1952.10.30.66. PL 20, fig. 56. Unio egyptiacus Cail. Egypt and Senegal. BMNH 1907.10.28.173. PL 20, fig. 57. Unio collinus Conrad. Virginia. BMNH 1907.12.30.60. PL 20, fig. 58. Unio subrotundus Lea. Ohio. BMNH 1907.10.28.195. PL 20, fig. 59. Unio ravenelianus Lea. Tennessee. BMNH 1907.12.30.61. PL 21, fig. i. Unio cooperianus Lea. Ohio. BMNH 1907.10.28.159. PL 21, fig. 2. Unio orbiculatus Hildreth. U. States. BMNH 1907.10.28.194. PL 21, fig. 3. Unio paranensis Lea. R. Parana. BMNH 1907.10.28.161. PL 21, fig. 4. Unio lens Lea. Ohio. BMNH 1907.10.28.171. PL 21, fig. 5. Unio capsaeformis Lea. N. America. BMNH 1907.10.28.241. PL 21, fig. 7. Unio ellipsis Lea. Ohio. BMNH 1907.10.28.242. PL 21, fig. 8. Unio (Alasmondonta) rugosus Barnes. N. America. BMNH 1907.12.30.367. PL 21, fig. ii. Unio retusus Lamarck. N. America. BMNH 1952.10.30.59. PL 21, fig. 13. Unio littoralis Lamarck. France, &c. BMNH 1907.10.28.158. PL 21, fig. 14. Unio vancosus Lea. Ohio. BMNH 1907.12.30.36. PL 21, fig. 22. Unio pectorosus Conrad. Tennessee. BMNH 1952.10.30.67. PL 21, fig. 24. Unio verrucosus Barnes. Hab.? BMNH 1907.10.28.164. PL 21, fig. 25. Unio australis Lamarck. New Holland. BMNH 1952.10.30.64. PL 21, fig. 29. Unio auratus Lea. N. America. BMNH 1907.12.30.148. PL 21, fig. 30. Unio bilineatus Benson. R. Hoogly, Hindostan. BMNH 1952.10.30.63. PL 21, fig. 35. Unio triangularis Barnes. North America. BNMH 1907.10.28.250. PL 21, fig. 37. Unio iris Lea. Ohio. BMNH 1907.10.28.193. PL 23, fig. 8. Unio medellinus Lea. Vera Cruz. BMNH 1907.10.28.184. PL 23, fig. 55. Unio leeai Gray Ohio, BMNH 1907.10.28.249. PL 23, fig. 56. Unio spinosus Lea. Georgia in America. BMNH 1907.12.30.37. PL 23, fig. 59. Unio (Alasmodonta) bonellii Fe'russac. S. Germany. BMNH 1907.10.28.168. PL 23, fig. 60. Unio delphinulus Morelet. Yucatan. BMNH 1952.10.30.134. PL 24, fig. i. Unio foliatus Hildreth. United States. BMNH 1907.10.28.178. PL 24, fig. 2. Unio haysianus Lea. N. America. BMNH 1907.10.28.185. PL 24, fig. 3. Unio hembeli Conrad. Louisiana. BMNH 1907.12.30.35. PL 24, fig. 4. Unio cucumoides Lea. Australia. BMNH 1907.10.28.244. PL 24, fig. 5. Unio grayanus Lea. China. BMNH 1907.10.28.170. PL 24, fig. 8. Unio ventricosus Barnes. North America. BMNH 1907.10.28.256. PL 24, fig. 10. Unio (Margaritana) curreyanus Lea. Tennessee. BMNH 1907.10.28.266. UNIONACEA: TYPES AND FIGURED SPECIMENS 107 PL 24, fig. 13. Anodonta wahlamatensis Lea. Near the Rocky Mountains. BMNH 1907.12 30-33- PL 24, fig. 15. Anodonta angulata Lea. California. BMNH 1907.12.30.32. PL 24, fig. 17. Anodonta ensiformis Spix. Brazil. BMNH 1907.12.30.156. PL 24, fig. 18. Anodonta blainvilliana Lee. Chili? [on plate caption, spelled blainvilleana]. BMNH 1908.12.15.19. Specimens of Unionacea, exclusive of types, figured in : Hanley, S., 1863, Photographic Conch- ology, a second, or photographic series of the Conchological Miscellany & c. pp. 3 : 7 photo pis. col. 4to. London. No localities are given in this work, the localities are from the boxes in which the specimens were found. PL i, fig. 2. Anodonta ferruginea Lea. [Indiana]. BMNH 1908.12.15.15. PL i, fig. 3. Anodonta subcrassa Lea. [Philippine Is]. BMNH 1908.12.15.11. PL i, fig. 5. Anodonta ferussacciana Lea. [Ohio]. BMNH 1908.12.15.10. PL i, fig. 6. Anodonta cumingii Lea. BMNH 1908.12.15.1. PL i, fig. 9. A nodonta pavonia Lea. [Ohio]. BMNH 1908.12.15.17. PL 2, fig. i. Unio menziesii Gray. [New Zealand]. BMNH 1908.12.15.5. PL 2, fig. 2. Unio decurvatus Rossmassler [Germany]. BMNH 1908.12.15.3. PL 2, fig. 9. Unio trirostris. [Hindustan] BMNH 1907.12.30.45. PL 2, fig. 10. Unio prevostianus Lea. [Georgia]. BMNH 1908.12.15.18. PL 3, fig. 2. Unio lineatus Lea. [Georgia]. BMNH 1908.12.15.7. PL 3, fig. 4. Unio boykinianus Lea. [Alabama]. BMNH 1908.12.15.16. PL 4, fig. i. Unio bigbyensis Lea. [Tennessee]. BMNH 1908.12.15.2. PL 4, fig. 2. Unio crocodilorum Morelet. [Central America]. BMNH 1908.12.15.4. PL 4, fig. 5. Unio preovalis Conrad. [Alabama]. BMNH 1908.12.15.6. PL 4, fig. 8. Unio spatulatus Lea. [Wisconsin]. BMNH 1908.12.15.14. PL 5, fig. i. Anodonta rubens Lamarck. [Lower Nile]. BMNH 1098.12.15.8. PL 5, fig. 2. Mycetopus siliquosus Spix. [Panama River]. BMNH 1908.12.15.12. PL 5, fig. 4. Unio (Symphonata) cumingii Lea. [Alabama]. BMNH 1908.12.15.9. Specimen of Unionidae figured in ; Forbes, E. and S. Hanley, 1853, A History of British Mollusca and Their Shells. PL 40, fig. 3. Anodonta cygnea Linnaeus. BMNH 1907.12.30.53. Specimens of Unionacea, exclusive of types, figured in : Hanley, S. and W. Theobald, 1870-76. Conchologia Indica ; Illustrations of the Land and Freshwater Shells of British India. PL 9, fig. 3. Trigonodon crebristriatum Anthony. British Birmah. BMNH 1907.10.28.204. PL 9, fig. 4. Pseudodon salwenianum Gould. Salwen River, Birmah. BMNH 1907.12.30.44. PL 9, fig. 5. Trigodonodon crebristriatus, var. Anthony. Hab.? BMNH 1907.10.28.203. PL 9, fig. 6. Unio lamellatus [generosus on box], var. Lea. A perculiar winged form from Mandelay. BMNH 1907.10.28.254. PL 10, fig. i. Unio olivarius Lea. Rohilcund Streams. BMNH 1907.12.30.62. PL 10, fig. 4. Unio macilentus Benson. " Bengal ", Mandelay. BMNH 1907.12.30.41 PL 10, fig. 6. Unio bonneaudi Eydoux and Souleyet. Bhama, [Bhamdo] Upper Birmah (Blan- ford). BMNH *4 15.06.1.1. PL 10, fig. 7. Unio pugio Benson. Ava. Pegu (Theobald). BMNH 1907.10.28.258. PL ii, fig. la. Unio favidens Benson. Sunderbund, Bengal. BMNH 1907.12.30.43. PL n, fig. 5. Unio crispisulcatus Benson. Tenasserim ; Pegu. BMNH 1907.10.28.263. PL ii, fig. 9. Unio trirostris Benson, non Reeve 1868. Moradabad. BMNH 1907.12.30.47. PL 12, fig. 2. Unio gerbidoni Eydoux and Souleyet. " Coromandel ". BMNH *4i8.o6.i.i. PL 12, fig. 3, 3a. Unio caeruleus Lea. River Hooghly, &c. BMNH 1907.12.30.63. PL 12, fig. 4. Unio gerbidoni, var. Eydoux and Souleyet. Hab.? BMNH 1907.12.30.67. * Goodwin-Austen's catalogue number. io8 R. I. JOHNSON PI. 12, fig. 6a. Unio leioma Benson. Near Bombay. BMNH 1907.12.30.66. PL 42, fig. i. Unio birmanus Blanford. Bhamo, Upper Birmah. BMNH *55i.o6.i.i. PL 42, fig. 3. Unio foliaceus Gould. Tavoy, Birmah, Pegu. BMNH 1907.10.28.240. PL 42, figs. 5, 6. Unio marcens Hanley (for U.favidens, var. marcens of Benson). Berhampooter River, Assam. BMNH *424.o6.i.i. PL 42, fig. 7. Unio marginalis, var. anodontina Kuster (U. anodontinus, Kiister non Lamarck). River Godavery ; Nagpoor ; Sylhet. BMNH *45i.o6.i.i. PL 43, fig. 4. Unio marginals var. candaharica Hutton. River Sutlej. BMNH 1907.10.28.176. PL 44, fig. 4. Unio marginalis, var. corriana Lea (U. corrianus Lea). Near Calcutta &c. BMNH 1907.10.28.187. PL 44, figs. 5, 6. Unio corrugatus, var. laevirostris Benson (U. laevirostris Benson). River Godavery ; Pemguang &c. BMNH 1907.10.28.260. PL 45, fig. 2, Unio corrugatus Miiller, var. solida. From the River Godavery. BMNH *42O. 06.1.1. PL 45, fig. 3. Unio corrugatus Lea. Nagpoor ; Pemgunga. BMNH *6o3.o6.i.i. PL 45, fig. 6. Unio wynegungaensis Lea. River Wynegunga, &c. BMNH 1907.12.30.58. PL 46. fig. 3. Unio scobinal, var. Benson. Belgaum, Deccan. BMNH *5i5.o6.i.i. PL 107, fig. 2. Unio triembolus Benson. Nerbudda River. BMNH *493.o6.i.i. PL 107, fig. 4. Unio indicus, var. aurea Sowerby. Nerbudda River. BMNH 1907.12.30.40. PL 107, figs. 6, 7. Unio sikkimensis Lea. Assam. BMNH *24i6.O3.viii.i. PL 154, fig. 2. Unio parma ?, var. Benson. Bhamao. BMNH *6i7.o6.i.i. PL 154, fig. 3. Unio rugosus Gmelin. Coromandel. BMNH 1907.12.30.64. PL 154, fig. 4. Unio mandely ay a nus Theobald. Mandelay, Birmah. BMNH 1907.12.30.49. PL 154, fig. 5. Unio macilentis, var. Surat. Near Chimoor ; Pern Gunga. BMNH *622.o6. i.i. PL 154, fig. 7. Unio tavoyensisl , var. ; Birmah. BMNH 1907.12.30.376. PL 155, fig. 2. Unio bhamaoensis Theobald. Near Bhamae, and from Western Prome, Pegu. BMNH 1907.10.28.169. * Goodwin- Austen's catalogue number. R. I. JOHNSON MUSEUM OF COMPARATIVE ZOOLOGY HARVARD UNIVERSITY CAMBRIDGE, MASS. 02138, U.S.A. PLATE i FIG. i. Iridina welwitschii Morelet. La riviere Muria, pres de Trombeta (Golungo-Alto) Angola. Lectotype BMNH 93.2.4.1740. Length 86, height 48, width 22 mm. FIG. 2. Anodon lucasii Morelet. La Calle [Algeria]. Holotype BMNH 93.2.4.1950. Length 107, height 56, width 33 mm. FIG. 3. Spatha baikii A. Adams. River Niger [Nigeria]. Lectotype BMNH 196466. Length 116, height 75, width 46 mm. Bull. Br. Mus. nat. Hist. (Zool.) 20, 3 PLATE r PLATE 2 FIG. i. Unio episcopalis Tristram. Orontes River, Palestine. Lectotype BMNH 1936.3.10.3 Length 80, height 47, width 28 mm. FIG. 2. Unio simonis Tristram. Orontes River, Palestine. Holotype BMNH 1936.3.10.6. Length 48, height 35, width 24 mm. FIG. 3. Unio scamnatus Morelet. Rio Tacataco [Pinar del Rio] Cuba. Lectotype BMNH 1893. 2.4.1976. Length 52, height 34, width 17 mm. FIG. 4. Anodonta tunizana Morelet. (La Calle), Tuniza. Lectotype BMNH 1893.2.4.1964. Length 55, height 32, width 20 mm. FIG. 5. Unio calamitarum Morelet. Rivulum Baluntie, propre Palenqueanum vicum [Chiapas, Mexico]. Lectotype BMNH 1893.2.4.2010. Length 54, height 32, width 26 mm. FIG. 6. Unio sitifensis Morelet. L'oued dehhab, pres d'Bone, Algeria. BMNH 1893.2.4.1965. Length 71, height 33, width 23 mm. Bull. BY. Mus. nat. Hist. (Zool.) 20, 3 PLATE 2 Printed in Great Britain by Alden & Mowbray Ltd at the Alden Press, Oxford J A REVIEW OF THE SPECIES OF HEMILEPISTUS S.STR. BUDDE-LUND, 1885 (ISOPODA, PORCELLIONIDAE) R. J. LINCOLN BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 20 No. 4 LONDON : 1970 A REVIEW OF THE SPECIES OF HEMILEPISTUS S. STR. BUDDE-LUND, 1885 (ISOPODA, PORCELLIONIDAE) v BY ROGER JOHN LINCOLN Pp. 109-130 ; 8 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 20 No. 4 LONDON : 1970 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, is issued in five series corresponding to the Departments of the Museum, and an Historical series. Parts will appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department. This paper is Vol. 20, No. 4 of the Zoological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation Bull. Br. Mus. nat. Hist. (Zool.). Trustees of the British Museum (Natural History), 1970 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 10 December, 1970 Price 80 p A REVIEW OF THE SPECIES OF HEMILEPISTUS S. STR. BUDDE-LUND, 1885 (ISOPODA, PORCELLIONIDAE) By R. J. LINCOLN SYNOPSIS A brief historical review of relevant literature is given. At the present time the genus Hemi- lepistus is divided into two sub-genera, Desertellio and Hemilepistus sensu-stricto. The latter group is dealt with in this paper. The sub-genus Hemilepistus contains nine recognised species, with a range of distribution from North Africa, through the Near Eastern countries, into central Asia. A diagnosis, with figures, is given for each species, together with a key for their identi- fication. Details of distribution are provided in each case. The synonymy for each species is presented. Where possible type material has been studied, and the locations of the types are indicated in the lists of material examined. INTRODUCTION BUDDE-LUND (1879) proposed the division of the genus Porcellio Latreille into seven sub-genera, but did not give any descriptive details of these divisions until the publication of his monograph on terrestrial isopods in 1885. As erected by Budde- Lund, the sub-genus Hemilepistus contained 10 species, of which 4 were newly described. The earliest description of a terrestrial isopod which can be attributed to Hemi- lepistus is found in an account of a journey through Russia by Pallas (1771). Further species are contained in the beautifully illustrated account of the fauna of Egypt com- piled by Audouin and Savigny (1826), and amongst the crustacean fauna of Turkestan described by Uljanin (1875). Brandt (1833) and Milne-Edwards (1840) published lists of the species of Porcellio known at that time, and included a number of new species which were later placed under Hemilepistus. Budde-Lund (1885) split the species of Hemilepistus into two groups, each of 5 species, on the basis of the presence or absence of a median frontal line between the frons and the epistome. Verhoeff (1930) used this character to erect two new sub- genera, Hemilepistus and Desertellio. This raised Hemilepistus Budde-Lund to generic status, a move which appears to have been adopted much earlier by Budde-Lund and Stebbing (1911). In a subsequent account of isopods from the Mediterranean coasts, Verhoeff (1931) describes two new species of Hemilepistus, but he makes no mention of the sub-genera which had been proposed. Arcangeli (1932) considers a number of the species of Hemilepistus to be invalid and arrives at a lengthy synonymy under crenulatus Pallas. More recently some Russian zoologists have made studies of this group of terrestrial isopods with a special interest in their biology and ecology. Borutzky (1945) gives a description of the woodlice fauna of Turkmeniya and central Asia, and produces 112 R. J. LINCOLN the first key for identification of the species of that area. He adopts the sub-genera of Verhoeff, and adds a further five new species. In a subsequent paper on Hemi- lepistus, Borutzky (1958) gives another key and drawings of a number of species. However, this account differs in many respects from the earlier paper and indicates Borutzky's uncertainty about the validity of some members of the genus. In undertaking this review of the sub-genus Hemilepistus s. str. the type material has been examined wherever possible. Much of the Budde-Lund collection is held in the British Museum (Natural History), together with a number of the Verhoeff syntypes. The location of other type material is given in the relevant parts of the text. Hemilepistus Budde-Lund, 1879 Body long, convex, grey or greyish-brown in colour with lighter tuberosity ; head and anterior peraeon tergites with armature of conical tubercles, which may be small and rounded or developed into large prominent crests ; lateral lobes on head small and set at an oblique angle ; eyes large, convex ; median frontal lobe on head small or absent ; pleon small, narrower than peraeon, smooth dorsally ; telson triangular with rounded apex ; antenna 2 short, with strongly developed peduncle and small, two segmented flagellum ; exopods of pleopods large, expanded ; 2-5 prs of pseudo- tracheae. KEY TO THE SUB-GENERA OF Hemilepistus 1 Frontal median lobe of head absent ...... Hemilepistus s. str. 2 Frontal median lobe present, either entire or cleaved in the centre Desertellio Verhoeff Hemilepistus (Hemilepistus) Budde-Lund, 1879 Body elongate, slate grey or brown with tubercles lighter in colour ; head with well defined pattern of conical tubercles on dorsal side, sometimes a number of additional smaller tubercles within basic arrangement ; lateral lobes of head oblique with small projecting upper lobes ; median lobe and suture line absent ; peraeon tergites 1-3 with well developed tubercles along posterior and lateral margins ; tubercles may form prominent crests ; (arrangement of tubercles on lateral and posterior margins of tergites 1-3 is expressed numerically as the " Dental formula ") ; peraeon tergites 4-7 smooth ; peraeon tergite i with prominent antero-lateral projection of epimera, apex pointed or acutely rounded ; pleon short, smooth, and with pointed, back- wardly directed, epimera ; telson triangular, usually with slight dorsal depression, margins either straight or concave ; antenna i very small ; antenna 2 short, reaching only as far as posterior edge of peraeon tergite i, peduncle strongly developed, segment 2 usually with prominent inner lobe, segment 5 elongate ; flagellum markedly shorter than last segment of peduncle ; flagellum segments either sub-equal, or segment i longer than segment 2 ; mandible large, 4-7 penicilli (fig. ik) ; maxilla 2 small, bilobed (fig. il) ; maxilla i and maxillipeds strongly developed (figs. li, j] ; peraeopod i, <$ (fig. ib) small, basis long, merusand carpus with long spines ; peraeopod REVIEW OF HEMILEPISTUS B-L. 113 7, 6 $ $, length 14-19 mm, width 5-5-7-5 mm. Collected in the Caucasus. Leningrad Museum. Cat. no. 1677. DIAGNOSIS. Body broad, length 13-20 mm, width 4-5-7-5 mm, greyish-brown with tubercles and epimera paler in colour ; peraeon somewhat rectangular, tergites 5-7 slightly broader than rest ; head with 16-20 rounded tubercles in a character- istic pattern over a triangular area ; pattern consists of large circle in middle of head with lateral rows of 3-4 tubercles extending to postero-lateral corners (fig. 2c, d) ; sometimes 1-2 tubercles in centre of the large circle and a number of smaller tubercles within the triangular area ; head with prominent lateral lobes, upper part of lobes rounded, outer edges sinuous (fig. 2a) ; ratio of width of head to width of peraeon tergite 2 is about i : 1-8 ; peraeon tergite i, antero-lateral projection of epimera short, only slightly upturned and with rounded apex (fig. 20) ; peraeon tergites 1-3 with short, conical tubercles along posterior and lateral margins ; outer tubercles along posterior margin more bulbous than inner ones, those on tergite i set at right angles to dorsal surface ; posterior edge of tergite i strongly developed ; tubercles on tergites 2-3 projecting backwards parallel to dorsal surface ; peraeon lateral tubercles prominent, anterior tubercle on tergite i largest, flattened and rectangular ; tergite 4 with only faint traces of tuberosity ; tergites 5-7 smooth ; (dental formula, tergite i, 3-4, 14, 3-4 ; tergites 2 & 3, 2-3, 14, 2-3) ; in large specimens tubercles on tergites 1-3 may form prominent crests ; telson triangular with shallow dorsal depression, lateral margins concave (fig. ze) ; antenna 2, peduncle segment 2 with large inner lobe, flagellum about f length of peduncle segment 5, flagellum segment i longer than 2 (fig. 26) ; mandibles (fig. i&), maxillules (fig. i/), maxillae (fig. il), maxillipeds (fig. li), peraeopods i (fig. la), and peraeopods 2 (fig. 16), as figured ; exopods of pleopods 1-5 (£ (fig. ic-g), pseudotracheae on pleopods 1-2 very well developed, small on 3 and rudimentary on 4-5 ; exopods of pleopod i £, with strongly sinuous posterior margin and prominent rounded inner angle (fig. ic). DISTRIBUTION. Caucasus ; Apsheron peninsular ; northern Iran. REMARKS. Budde-Lund (1885) gives two separate localities for the distribution of klugii. The first is " Caucasus " from material in the Berlin Museum, and the second is " Schiras in Persia " based upon three specimens in the Copenhagen Museum. Both of these collections have been examined and it is clear that they represent quite different species. The material from the region of the Caucasus is the true klugii. The other specimens from Iran will be described later as a new species. Borutzky (1958) was the first to place doubt on the validity of the Budde- Lund klugii from Schiras and included it in a list of species of uncertain taxonomic position, but without giving any reasons. 2. Hemilepistus (H.) crenulatus (Pallas, 1771) (text-figs. 3a-h) Oniscus crenulatus Pallas, 1771 : 477. Porcellio crenulatus ; Latreille, 1804 : 46. REVIEW OF HEMILEPISTUS B-L. 117 Porcellio elegans Uljanin, 1875 : 6. Hemilepistus elegans ; Budde-Lund, 1879 : 4 ; 1885 : 154 ; Borutzky, 1945 : 195. Hemilepistus crenulatus ; Budde-Lund, 1885 : 153 ; Arcangeli, 1932 (part) : i ; Borutzky, 1958 : 1465. Hemilepistus pectinatus Budde-Lund, 1885 : 153 ; Borutzky, 1945 : 195 ; 1958 : 1467. MATERIAL EXAMINED. 2 $ $, length 14 mm, width 4-5 mm. Syntypes of elegans Uljanin, collected in valley of Syr-darya, Turkestan. Berlin Museum, cat. no. 6630 (figs. 36, d, g). i (£, (i juvenile), length 16-5 mm, width 5-0 mm, collected by Dr. Pawlowsky in Turkestan. B.M. (N.H), (figs. 30, c, e,f), reg. no. 1916.12.2, 3-4. i $, (damaged) collected at Schrenck, Kazakhstan. Leningrad Museum, cat. no. 1737- 1 $, length 15-0 mm, width 5-0 mm. Holotype of pectinatus Budde-Lund, col- lected at Schrenck, Kazakhstan. Leningrad Museum, cat. no. 1769. 2 <£<£, length 14-15 mm, width 4-5-5-0 mm. Budde-Lund collection from Turkestan. B.M. (N.H.), reg. no. 1921.10.18, 4095-4096. DIAGNOSIS. Body long and narrow, length 14-16-5 mm, width 4-5-5-0 mm ; colour dark grey with pale yellow tubercles. Size of tubercles varies considerably with body size, figs. 30, c, e, f are of large male specimen with strong tuberosity ; figs. 3&, d, g of smaller specimen with weak tuberosity ; head with 10-12 tubercles, 6 forming a semi-circle in the middle with 2-3 in rows extending towards postero-lateral corners of head (figs, y, d) ; very few additional tubercles developed on head ; on large specimens head-tubercles strong, conical and pointed ; smaller specimens with weak, rounded tubercles ; lateral lobes of head large, outer edge straight or slightly convex, upper lobe somewhat pointed (fig. 30) ; ratio of width of head to width of peraeon tergite 2 is about i : 1-6 ; peraeon tergite i, antero-lateral projection of epimera short, pointed and with sinuous lateral margin ; tergites 1-2 on specimen with strong tuberosity with large, conical, pointed tubercles along posterior and lateral margins (fig. 30) ; prominent crests developed ; tergite 3 with small bulbous tubercles, tergite 4 with small laterals only, tergites 5-7 smooth ; on small specimens with weak tuberosity tergites 1-2 with small rounded tubercles (fig. 36), no crests developed, tergites 3-7 smooth ; (dental formula, i, 4-5, 14, 4-5 ; 2, 3-4, 14, 3-4 ; 3, 2-3, 12, 2-3) ; small specimens have only 12 posterior marginal tubercles on tergites 1-3 ; telson triangular with shallow dorsal depression, lateral margins con- cave (3g,f) ; telson more pointed in larger specimens ; antenna 2, peduncle segment 2 with small inner lobe ; flagellum about f length of peduncle segment 5, flagellum segments sub-equal (fig. 30) ; exopods of pleopod i <$, only weakly sinuous posterior margin, inner angle not pronounced as in klugii (fig. ic) ; pseudotracheae well developed on exopods 1-2, small on 3, rudimentary on 4-5. DISTRIBUTION. Central Asia ; southern Kazakhstan, shores of Aral sea, valley of Syr-darya, region of Golodnaya Steppe, shores of river Ili, Kum-basy mountains ; crenulatus type locality in arid hills around lake Inder. n8 R. J. LINCOLN FIG. 3. Hemilepistus crenulatus (Pallas) ; a, lateral view of head and tergites 1-2 (large specimen) ; b, lateral view of head and tergite i (small specimen) ; c, dorsal view of head (large specimen) ; d, dorsal view of head (small specimen) ; e, antenna 2 ; /, telson (large specimen) ; g, telson (small specimen) ; bar scale i mm. 3. Hemilepistus (H.) reaumuri (Audouin & Savigny, 1826) (text-figs, ^a-d) Porcellio Reaumurii Audouin & Savigny, 1826 : 13 ; Milne-Edwards, 1840 : 170. Porcellio clairvilli Brandt, 1833 : 179. Porcellio syriaceus Koch, 1847. Hemilepistus Reaumurii ; Budde-Lund, 1879 : 4 ; 1885 : 155. REVIEW OF HEMILEPISTUS B-L. 119 Hemilepistus reamurii ; Dollfus, 1892 : 10 ; 1894 : 3 '• 1896 : 546 ; Richardson-Searle, 1926 : 206 ; Cloudsley-Thomson, 1955 : 248 ; Borutzky : 1958, 1471. Paraniamba tuber culata Collinge, 1914 : 206 Hemilepistus palaestinus Verhoeff, 1931 : 38. Hemilepistus bodenheimeri Verhoeff, 1931: 40. MATERIAL EXAMINED. 18 (J, length 15-22 mm, width 5-5-8-0 mm. Budde-Lund collection from various localities ; Tunisia, Cyrenaica, Algeria (Biskra and Algiers). B.M. (N.H.). 49 width 6-5-8-0 mm. Collection from Algeria (Biskra). Copenhagen Musuem. DIAGNOSIS. Body broad, length 11-23 rnm, width 4-0-8-0 mm, slate grey in colour with lighter grey epimera and whitish tubercles ; head with a large number (25-30) of small, pointed tubercles in a characteristic pattern ; pattern consists of large circle in middle of head, a group of tubercles in postero-lateral corners, and 8-10 tubercles in transverse row along posterior margin (fig. 46) ; lateral lobes of head prominent, upper lobe rounded, outer edge straight or convex (fig. 40) ; ratio of width of head to width of peraeon tergite 2 is about i : i -8 ; peraeon tergites 1-3 with large number of small tubercles, never developed into crest ; tubercles in middle of tergites pointed, lateral group somewhat rounded and flattened ; peraeon tergite i with posterior marginal row of tubercles, 4 tubercles in transverse median row, and 2 anterior marginal tubercles ; tuberosity on tergites 2-3 similar to that of tergite i except no anterior marginal tubercles ; tergite 4 with weak tuberosity ; sometimes faint traces of tubercles on tergites 5-7 ; (dental formula, i, 10-15, I4~I6, 10-15 > 2, 8-12, 14-16, 8-12 ; 3, 5-8, 14-16, 5-8) ; all tubercles small and number very variable ; telson triangular at base, deep dorsal depression, margins concave, apex acutely rounded (fig. 4^) ; antenna 2, peduncle segment 2 with very large inner lobe ; 120 R. J. LINCOLN flagellum half length of peduncle segment 5 ; flagellum segment i markedly longer than 2 (fig. 40) ; exopods of pleopod i $, sinuous posterior edge and prominent inner angle as in klugii (fig. ic) ; pleopods pigmented ; pseudotracheae 1-2 large, 3-5 rudimentary. FIG. 4. Hemilepistus reaumuri (Audouin & Savigny) ; a, lateral view of head and tergite i ; b, dorsal view of head ; c, antenna 2 ; d, telson ; bar scale i mm. DISTRIBUTION. Widely spread through Syria, Palestine, Egypt, Libya, Tunisia and western Algeria. According to Vandel (1955) it rarely occurs west of the meridian of Algiers. REMARKS. Verhoeff (1931) describes two new species of Hemilepistus from the neighbourhood of Jerusalem — palaestinus and bodenheimeri. The diagnosis of palaestinus was based upon the stronger armature of tubercles on the head and anterior three peraeon tergites, and the more spinose nature of the tubercles. Syntype material from the Verhoeff Collection in Munich Museum, and from the British Museum (Natural History), was examined together with material collected in many localities in North Africa, Israel and Syria. The degree of development of the tubercles is very variable and as a result of this work palaestinus and reaumurii are REVIEW OF HEMILEPISTUS B-L. 121 considered to be a single species. The bodenheimeri type specimens are small in size, a greyish brown colour, with white epimera. The tuberosity on the head and first three peraeon tergites is similar to reaumuri but only weakly developed. The telson is more triangular in shape than the telson of adult reaumuri, although it resembles the telson of small and juvenile reaumuri. A triangular telson is typical of the young stages of a number of species of Hemilepistus. The above features suggest that the bodenheimeri specimens are in fact small individuals of reaumuri. Without additional data concerning their distribution they are considered as a single species. 4. Hemilepistus (H.) cristatus Budde-Lund, 1879 (text figs. 5a-h) Porcellio klugii ; Lessona, 1867 (not Brandt) : 187. Hemilepistus cristatus Budde-Lund, 1879 : 4 ; 1885 : 153 ; Borutzky, 1945 : 193 ; 1958 : 1467. Hemilepistus elegans ; Walter, 1889 : mo. Hemilepistus crenulatus ; Arcangeli, 1932 (part) : i. IHemilepistus uljanini Borutzky, 1955 : 216 ; 1958 : 1469. MATERIAL EXAMINED, i $, i $, length 17 mm, width 5-0 mm. Syntypes cristatus Budde-Lund, collected from " Serdscen in Persia ". B.M. (N.H.), reg. no. 1956. 10.10, 156-157. 1 <$, 3 $$, length 17-18 mm, width 4-5-5-0 mm. Norman collection from " Serdscen in Persia ". B.M. (N.H.), reg. no. 10443, 46. 2 $<$, 12 ? ?, length 15-19 mm, width 4-5-6-0 mm. No locality. B.M. (N.H.), reg. no. 1970, 197. DIAGNOSIS. Body long and narrow, length 15-17 mm, width 4-5-6-0 mm, dark grey body, tubercles pale yellow ; head with 16-20 short conical tubercles in characteristic pattern within a triangular area ; pattern consists of a large circle of 8 tubercles in middle of head, with lateral rows of 3-4 tubercles extending to postero- lateral corners (fig. 56) ; sometimes 2-3 tubercles in centre of circle, and a number of smaller tubercles within the triangular area ; head tuberosity rather variable (figs. 5c-/), and in an extreme case the pattern is not apparent because the tubercles are poorly developed and flattened (fig. 5/) ; lateral lobes of head prominent, with rounded apex and sinuous outer edge (fig. $a) ; ratio of width of head to width of peraeon tergite 2 is about i : 1-5 ; peraeon tergite i, antero-lateral projection of epimera long, pointed and upturned at apex (fig. 5«) ; peraeon tergites 1-3 with well developed tubercles which may form tall upright crests on tergites 1-2 ; tergite 4 with weak tuberosity, best developed laterally ; tergites 5-7 smooth ; most anterior tubercle of tergite i largest, rectangular ; (dental formula i, 4, 14, 4 ; 2 & j, 3, 12, 3) ; telson triangular, shallow dorsal depression, lateral margins deeply concave, apex acutely rounded (fig. 5g) ; antenna 2, peduncle segment 2 with large inner lobe ; flagellum half length of peduncle segment 5 ; flagellum segments sub-equal or nearly so (fig. 5&) ; exopods of pleopod i <$, strongly sinuous posterior margin, inner angle broad and rounded but less prominent than klugii (fig. ic) ; pseudotracheae on 1-2 large, 3 small, 4-5 rudimentar}^. R. J. LINCOLN FIG. 5. Hemilepistus cristatus Budde-Lund ; a, lateral view of head and tergite i (Syntype) b, dorsal view of head (Syntype) ; c-f, dorsal view of head ; g, telson (Syntype) ; h, antenna 2 (Syntype) ; bar scale i mm. REVIEW OF HEMILEPISTUS B-L. 123 DISTRIBUTION. Iran ; central Asia, slopes of Kopet-Daga from Serakhs to Kazandzhik, valley of Sumbar Uzboy ; Ashkhabad ; type locality Serdscen in Iran. REMARKS. Borutzky (1955) describes a new species from Turkmeniya — uljanini. This seems to differ from cristatus only in the detailed tuberosity of the head. How- ever, it is clear that there is considerable variation in the size and arrangement of tubercles on the head of cristatus and the description of uljanini falls within this range. In other features given uljanini and cristatus appear to belong to the same species. 5. Hemilepistus (H.) magnus Borutzky, 1945 (text-figs. 6a-d) Hemilepistus (H.) magnus Borutzky, 1958 : 1467. MATERIAL EXAMINED, i <£, length 26 mm, width 10-5 mm, Budde-Lund collection from Turkmenistan. B.M. (N.H.), reg. no. 1921.10.18, 4146. DIAGNOSIS. Largest body size for species of Hemilepistus, length 26-30 mm, width 9-0-10-5 mm, peraeon nearly uniform in width, tergites 5-7 a little broader than rest ; colour grey ; according to Borutzky (1958) the ventral surface is dark grey with yellow spots ; head with 12-14 long, slender tubercles in characteristic pattern (fig. 6b) ; 6 largest tubercles form an open semi-circle on front of head, with rows of 3-4 tubercles extending to postero-lateral corners ; all tubercles on head very long, slender and rounded at apex ; lateral lobes of head small ; upper part of lateral lobe rounded outer margin concave (fig. 6a) ; ratio of width of head to width of peraeon tergite 2 is about 1:2-0; peraeon tergite i, antero-lateral projection of epimera short, rounded ; postero-lateral angle of epimera on tergite i forming an acute, backwardly pointing, process (fig. 6 a) ; peraeon tergites 1-3 with very long tubercles along posterior and lateral margins ; all tubercles long, slender, cylindrical, and well spaced apart ; tergite 4 with weak tuberosity, tergites 5-7 with traces of tuberosity, best developed laterally ; epimera of tergites 6-7 markedly swollen ; (dental formula j & 2 & 3, 3-5, 12-13, 3-5) ; pleon short and broad ; epimera of pleon long, pointed and curved upwards a little at apex ; telson wide at base, long and with acutely rounded apex, margins sinuous (fig. 6d) ; dorsal surface of telson flat or very weakly concave ; antenna 2 strongly developed, peduncle segment 2 with small inner lobe, flagellum half length of peduncle segment 5 ; flagellum slender, segment 2 half length of i (fig. 6c) ; exopods of pleopod i <£, sinuous posterior margin and rounded inner angle, but less pronounced than klugii (fig. 2c) ; pleopods pigmented. DISTRIBUTION. Turkmeniya, Fergana valley and the region of the Alayli moun- tains. 6. Hemilepistus (H.) reductus Borutzky, 1945 Hemilepistus (H.) reductus Borutzky, 1945 : 495 ; 1958 : 1468. MATERIAL EXAMINED. None. DIAGNOSIS. Body small, elongate, length 13-16 mm, width 4-5-5-0 mm ; dark I24 R. J. LINCOLN FIG. 6. Hemilepistus magnus Borutzky ; a, lateral view of head and tergite i ; b, dorsal view of head ; c, antenna 2 ; d, telson ; bar scale i mm. grey with lighter epimeral margins, tubercles whitish in colour ; head with about 12 small tubercles arranged in two curved rows extending from poster-lateral corners of head to an apex at front ; additional smaller turbercles may be present on the head ; lateral lobes of head short, upper edge straight ; ratio of width of head to width of peraeon tergite 2 is about 1:1-5 ; tubercles present on posterior and lateral margins of peraeon tergites 1-2 ; tergite i with well developed laterals, but posterior row very much reduced in the middle of the tergite ; tubercles on peraeon tergite 2 larger than on tergite i, uniform in size and closely set together ; tergite 3 with only a trace of lateral tuberosity, posterior margin smooth ; tergites 4-7 smooth ; (dental formula j, 4-7, 12-16, 4-7 ; 2, 4-7, 10-14, 4-7) ; antenna 2, peduncle segment 5 a little longer than flagellum ; flagellum segments sub-equal ; exopods of pleopod i with less sinuous posterior margin than klugii (fig. 20), and no lobe at inner angle. REVIEW OF HEMILEPISTUS B-L. 125 DISTRIBUTION. Kazakhstan ; Kumak, Kara-darya, the environs of the towns of Katta-Kurgan ; widely distributed in the valley of the river Zeravshan. 7. Hemilepistus (H.) rhinoceros Borutzky, 1958 Hemilepistus (H.) rhinoceros Borutzky, 1958 : 1469, fig. 8. MATERIAL EXAMINED. None. DIAGNOSIS. Body small, elongate, length 13 mm, width 4-0-4-5 mm, body grey, tubercles white, epimeral margins light grey in colour ; head with a single large tubercle situated in a median position towards the front ; this tubercle is divided on posterior side into two smaller tubercles ; small group of 3-4 tubercles above the eyes ; lateral lobes of head small with rounded upper lobe ; ratio of width of head to width of peraeon tergite 2 is about 1:1-3; peraeon tergites 1-2 with small conical tubercles directed upwards on tergite i and backwards on tergite 2 ; tergites 3-7 smooth, without tuberosity ; (dental formula i & 2, 4-5, 12, 4-5) ; telson triangular, apex pointed and lateral margins straight ; antenna 2, peduncle segment 5 one to one and a half times length of flagelrum ; flagellum segments sub-equal. DISTRIBUTION. Kazakhstan ; type locality Dzhusandala near Lake Balkhash in an area of saline loess. 8. Hemilepistus (H.) aphganicus Borutzky, 1958 (text-figs, ja-g) Hemilepistus (H.) aphganicus Borutzky, 1958 : 1470. Hemilepistus (H.) aphganicus kabulensis Borutzky, 1958 : 1471. MATERIAL EXAMINED. 5 ^^,3 $?, length 15-21 mm, width 5-0-7-0 mm. Collected by the Afghanistan Boundary Commission in the region around Bala Murghab, Afghanistan. B.M. (N.H.), reg. no. 86-50. 2 c?c?> 6 ??, (2 juveniles), length 19-20 mm, width 6-5-7-0 mm. Collected by the Afghanistan Boundary Commission from Serakhs, Turkmeniya, U.S.S.R. B.M. (N.H.), reg. no. 93.2.19, 1-12. DIAGNOSIS. Body broad, length 15-20 mm, width 5-0-7-0 mm, light brown in colour (in spirit) with pale yellow tubercles ; dry material grey ; peraeon rectangular, tergites 5-7 a little broader than rest ; head with 12-14 long, slender, rounded tubercles in a wide sinuous arc from postero-lateral corners towards the front (fig. 7&, c) ; 2-4 tubercles on centre of head, and row of 4-8 smaller tubercles along posterior margin ; arrangement of tubercles seen clearly in small individuals (fig. 76) ; large specimens may have a number of additional small tubercles on the head (fig. jc] ; lateral lobes of head with rounded apex, outer edge sinuous (fig. 70) ; ratio of width of head to width of peraeon tergite 2 is about i : 1-7 ; peraeon tergite i, antero-lateral projection of epimera long, apex rounded (fig. 70) ; peraeon tergites 1-3 with long, slender, rounded tubercles along posterior and lateral margins ; middle tubercles in posterior row somewhat smaller than others ; in large individuals, with strongly 126 R. J. LINCOLN developed tuberosity, tergites 1-2 have a tall crest of long, well spaced tubercles, those on tergite 2 larger than those on tergite i ; in small individuals, crests not developed, all tubercles small, equal in size, and directed backwards ; tergite 4 with small tubercles, tergites 5-7 smooth ; (dental formula J, 3-6, 14, 3-6 ; 2 & 3, 3-4, 14, 3-4) ; telson triangular, short (fig. ye) ; telson shape varies with body size ; figs. 7/ and 7g are taken from 15 mm and 10 mm specimens respectively ; at 10 mm stage, telson forms regular triangle, and at this stage head and peraeon tuberosity is just visible in characteristic pattern ; antenna 2, peduncle segment 2, with prominent FIG. 7. Hemilepistus aphganicus Borutzky ; a, lateral view of head and tergite i ; b-c, dorsal view of head ; d, antenna 2 ; e, telson (20 mm body length) ;/, telson (15 mm body length) ; g, telson (10 mm body length) ; bar scale i mm. REVIEW OF HEMILEPISTUS B-L. 127 inner lobe ; flagellum about half length of peduncle segment 5, flagellum segment i almost twice length of segment 2 (fig. yd] ; exopods of pleopod i $, with markedly sinuous posterior margin and prominent inner angle as in klugii (fig. ic) ; pseudo- tracheae on exopods 1-2 well developed, 3 small, 4-5 rudimentary. DISTRIBUTION. Afghanistan ; Turkmeniya, U.S.S.R. ; type locality around Yakatut in Afghanistan. REMARKS. Borutzky (1958) describes the species aphganicus from a small col- lection of dry material from Afghanistan. He also proposes a sub-species, kabulensis for a single female specimen from a locality near Kabul, although he adds that it may not be a valid sub-species because of the large variation between individuals of the species. The examination of the material in the British Museum (Natural History) does not justify the separation of the sub-species on the basis of the description given. 9. Hemilepistus (H.) schirasi n. sp. (text-figs. 8a-f) Hemilepistus klugii Budde-Lund, 1885 (not Brandt) : 152 (part). MATERIAL EXAMINED, i <£> i ?, length 15-17 mm, width 5-5-6-0 mm. $ Holo- type, $ paratype. Collected by Kollar from Shiraz in Iran. Copenhagen Museum. i <$, length 16 mm, width 5-5 mm. Paratype. CoUected by Kollar from Shiraz in Iran. B. M. (N.H.), reg. no. 1970 : 199. i bc ; cd lateral, dd : u nearly 0-5. Clitellum XIII or \ XIII-XX. Prostatic pores immediately lateral to setae b, on XVII and XIX, on round papillae, which are separated, longitudinally, by a space of equal width. Seta aoib sporadically absent in XVII and XIX ; b may be absent in XVIII. Male pores, not externally visible, on XVIII midway between the prostatic pores slightly lateral of b. Seminal grooves not recognizable. Spermathecal pores in 7/8 and 8/9 approximately midway between b and c lines. Pharyngeal glands ending in VII. Gizzard small, in VII, oesophageal musculature there considerably increased but diameter not greatly. Calciferous glands originat- ing posteriorly in IX ; lumen slitlike ; the very thick wall honeycombed by numerous blood spaces separated by stout trabeculae and confluent posteriorly to become fewer and larger. Last hearts in XI. Testes and funnels free in X. Seminal vesicles in IX and XI. Vasa deferentia not terminally thickened. Prostates passing gradually into muscular ducts about 200 ju long which discharge through cushion like thickenings ; terminal bursae absent. Spermathecae with ental portion tubular and twisted, with narrow lumen 6-8 \L or less in diameter ; further ectally a sharply demarcated portion with irregular cavity 20-28 /i in diameter and, finally, a short duct which has an extraordinarily thick muscular sheath ; diameter of spermatheca 34 \i in the ental tubular portion ; 70 [i in the ectal, swollen region. DISTRIBUTION. Paraguay : Makthlawaiya (mud of shallow pools after rain). THE OLIGOCHAETE GENUS EUKERRIA 147 REMARKS. The " cotypes " of this species in the British Museum (1930. 7. 30. 7/13) are all that the author has been able to trace. All are immature or lack the anterior and genital regions. Eukerria mcdonaldi (Eisen, 1893) Fig. gE-F Kerria mcdonaldi Eisen, 1893 : 294, PI. XI, fig. 1-6, 8-10, PL XII, fig. 13-27 ; Eisen, 1900 : 135 ; Michaelsen, 1900 : 372. ? Kerria zonalis Eisen, 1893 : 311, PI. XI, fig. 7, n, 12, PI. XII, fig. 28-30. Michaelsen, 1900 : 372. 1 = 25 mm, w = "i line ". All setae present in XVII-XIX, but setae ab here 1/3 smaller and slightly wider than other setae. Most setae with minute cicatricing at the free ends. Clitellum saddle-shaped, XIII-XX. Male genital field a raised area on each side of the ventral midline, separated by a cylindrical cavity crossing XVII-XIX ; this cavity bridged internally by arciform muscles. Prostatic pores paired in the setal zones of XVII and XIX shortly lateral of setae b ; male pores paired in XVIII, each on a small papilla, immediately lateral of setae b and therefore slightly median of the prostatic pores. Seminal groove on each side connecting the prostatic pores and curving slightly medially but deflecting a little laterally to skirt the male pore which it does not include. The prostatic pores lying on transversely oblong papillae which are thicker laterally. These papillae and the body wall medial of the seminal groove forming an approximately crescent shaped genital zone on each side. Female pores in front of setae ab of XIV. Spermathecal pores paired, in 7/8 and 8/9, or in 8/9 only, in cd lines. Last septal glands in VII ; oesophagus in \ IV to anterior XII, gizzard very rudi- mentary in VII. Calciferous glands arising from the oesophagus anteriorly in IX ; hidden by the oesophagus in dorsal view ; each rounded and blunt, with a single internal cavity with large projecting ridges and traversed by longitudinal blood vessels. Intestine commencing abruptly, anteriorly in XII. Gut highly vascular- ized in XI-XX. Hearts in X and XI. Nephridia commencing in IV ; with peritoneal cells in IX posteriorly ; ducts avesiculate. Testes and large sperm funnels in X ; unpaired (?) " sperm sacs " or " sperm masses " in X and XI ; seminal vesicles absent from IX. Vasa deferentia superficial on the parietes and very tortuous, ending at the male pores, in XVIII, without terminal dilatation. Excep- tionally with a second pair of sperm funnels in XI, and additional seminal vesicles in XII ; the anterior pair of sperm ducts opening adjoining the anterior prostatic pores in XVII, the posterior pair at the male pores in XVIII. Prostates 2 pairs (abnormally double on each side) much bent, when extended about as long as the width of a segment ; glandular part of the anterior and posterior prostates about three times and five times as long respectively as the muscular duct ; ducts of the anterior much narrower than those of the posterior pair ; neither with terminal expansion. Ovaries (palmate) and funnels in XIII ; ovisacs absent. Spermathecae with large, saclike ampulla and narrow tubelike duct, and usually, at their junction, with a diverticulum which is 3-lobed ; spermatozoa stored in the ampulla, not in the diverticula ; the 148 B. G. M. JAMIESON ampulla usually bent on the duct. The anterior spermathecae usually smaller, never larger, than the posterior pair. Spermatophores [?] paddle-shaped. DISTRIBUTION. Baja California : Miraflores near San Jose del Cabo (In mud) ; Cape Region (a pond near Santa Ana). REMARKS. E. zonalis agreed with E. mcdonaldi, with which the single specimen was collected, in lacking a gizzard, in the location of spermathecal, male and prostatic pores and in the possession of spermathecal diverticula and according to Eisen " much resembles " E. mcdonaldi. Differences from mcdonaldi were persistence of setae b in XVII and XIX, absence of spermathecae from VIII and duplication of the prostate glands on each side. On the whole resemblence to mcdonaldi is so close, even to the most unusual possession of spermathecal diverticula, and the chief difference, duplication of the prostate, is so clearly an abnormality that zonalis is here regarded as a junior synonym. Elsewhere in the Ocnerodrilinae spermathecal diverticula are seen only in Pygmaeo- drilus. There, however, they store sperm as is usual in the megascolecoids. The presence in E. mcdonaldi of " sperm sacs " in X and XI (Eisen, 1893) was subsequently denied (Eisen, 1900). In the latter account there were said to be only " sperm masses ", in X and XI. It seems likely that there were free sperm masses in X and seminal vesicles in XI. Eukerria papillifera (Rosa, 1895) Fig. 90 Kerria papillifera Rosa, 18953, : 3 ; Rosa, iSgsb : 145, PI. fig. 19-21 ; Michaelsen, 1900 : 370. 1 = 55-60 mm, w = 2 mm, s = 140. Epilobous. aa somewhat smaller than be. Setae ornamented distally by 4-5 longitudinal rows of arcuate depressions. Clitellum saddle-shaped, XIII — XIX, interrupted in aa and by the male field. Prostatic pores in ab lines on minute papillae in sucker-like depressions on large dome-shaped papillae. Seminal grooves absent. Male pores in XVIII in line with the prostatic pores. Ventral setae present on XVIII, absent from XVII and XIX. 3 unpaired midventral genital papillae in the posterior halves of XIV, XV and XVI. Sperm- athecal pores large, in setal lines b, with tumid lips. Gizzard absent. Calciferous glands round-based cones. Testes? Prostates tortuous or straight, extending through as many as 20 segments; ducts about 3 segments long each opening through a muscular copulatory sac. Posterior pair of spermathecae larger than the anterior ; ampulla oval, wider anteriorly, with short, wide, sharply demarcated duct, without diverticula. DISTRIBUTION. Central Paraguay. REMARKS. Two specimens in Torino Museum (ol. 113, ex. 289) collected by L. Borelli in Central Paraguay, are presumably syntypes but neither possesses the clitellar end. THE OLIGOCHAETE GENUS EUKERRIA 149 Eukerria pascuorum (Stephenson, 1931) Kerria pascuorum Stephenson, 1931 : 316. PI. fig. 10. 1 = 33-60 mm, w= 0-8 mm, s = (no?)-i44. Prolobous. Setae : aa = 3 ab — be ; ab — cd ; dd : u — 0-5. Clitellum saddle-shaped, XIV-XX and (sections) the greater part of XII. Prostatic pores on XVII and XIX in be, nearer b than c lines, on small porophores which are carried on moderately large, conspicuous papillae twice as long as wide, occupying the length of their segments and a little of XVIII ; a small part of XVIII equal to the diameter of a papilla intervening between the papillae of a side ; seminal grooves straight. Male pores (in sections) intermediate between and in line with the prostatic pores. Ventral surface of XVIII sometimes tumid. Spermathecal pores (from sections) in 7/8 and 8/9 in c lines. Last septal glands in VII. A moderate gizzard, with thick walls, but not much wider than the oesophagus, in VII. Calciferous glands arising posteriorly in IX ; pear-shaped with broad end anterior ; lumen slit-like or star-shaped, walls honey- combed as in limosa. Last hearts in XI. Testes and funnels free in X ; seminal vesicles in IX and XI lobulated or racemose. Prostates extending posteriorly through several segments ; duct equal in diameter to glandular part but muscular. Walls of glandular part one cell thick. Duct equal to or a little more than a segment in length, widening close to its termination at the surface of its porophore but lacking a terminal bursa or special muscular investment. Spermathecal ampulla elongated- ovoid or cylindrical ; sometimes bent on itself and sometimes constricted at the bend ; duct short but so muscular as to equal ampulla in width. DISTRIBUTION. Paraguay : Makthlawaiya (Mud of ponds in pasture). MATERIAL EXAMINED. 3 syntypes, immature or lacking the anterior and genital region, excluded from the above description ; B.M. (N.H.), 1930. 7. 30. 51/53. REMARKS. Distinctions from the sympatric E. limosa are few and of doubtful importance. The small lumen of each calciferous gland and the more ventral loca- tion of the Spermathecal pores appear to separate both species from the otherwise rather similar E. eiseniana. Eukerria rosae (Beddard, 1895) Fig. 4A-F Kerria rosae Beddard, 1895 : 224 ; Beddard, 1896 : 41 ; Michaelsen, 1900 : 372 ; Pickford, 1928 : 381, Fig. 5. 1 = 25-35 mm, w = 1-1-2 mm, s ?. Setae closely paired ; in segment XII aa \ab: be : cd : dd = 2-8 : i : 3-4 : 0-81 : 10-6 ;dd:u = 0-45 (i type-specimen, B. M. (N.H.)) ; setae a present on the male field, b present or absent. Nephropores conspicuous small papillae anteriorly in their segments about 1/3 be below c lines or (Hamburg material) not visible. Clitellum imperfectly developed. Prostate pores on small papillae nearer b than c, on XVII and XIX, each surrounded by a low, laterally elevated auricular lobe limited to its segment ; the 2 pores of a side connected by a seminal groove which is only slightly bent medianwards, and is bordered by slightly 150 B. G. M. JAMIESON tumid ridges. Male pores not externally apparent, from internal examination, in the seminal grooves at mid XVIII. Female pores on small circular papillae anterior in XIV, slightly lateral of b lines. Spermathecal pores inconspicuous, bordered anteriorly and posteriorly by slight ridges or on small papillae, about one-third be below setal lines c. Last septal glands in VII. Gizzard barely twice the width of the oesophagus but FIG. 4. A-F. Eukerria rosae, syntype, B.M. (N.H.), 1904.10.5. 929 : A and B, right anterior and posterior prostates ; C and D, left posterior and right anterior sperm- athecae respectively ; E, former spermatheca cleared (freehand) ; F, male genital field. G-H, syntype, Hamburg Museum, V. 4103 : G, left spermatheca of VIII ; H, male genital field of left side of same. I-L, E. saltensis, syntype, B.M. (N.H.), 1904.10.5 928 : I, Spermathecal pores ; J, prostates ; K, clitellar region ; L, left side of same. THE OLIGOCHAETE GENUS EUKERRIA 151 firm and thickly muscular. Calciferous glands almost sessile stoutly pear-shaped, broad end anterior, adpressed medianly below the oesophagus beyond which they project laterally. The walls of each pouch fairly thick, permeated by blood vessels, and projecting in places as folds into the lumen. Intestinal origin in XII ; oeso- phageal valve well developed at approximately £ XII. Typhlosole absent. Last hearts, in XI, exceedingly large, those in X less so. Nephridia avesiculate, the first in VII. Proandric ; sperm funnels multilocular, in X ; seminal vesicles in IX and XI; sperm ducts slightly widened in XI. Prostates extending posteriorly into XXIII. Glandular parts exceedingly long ; except ectally, thread-like and much coiled and mutually entangled ; maximally 90 ju wide. Ducts muscular and glossy, gently curved or strongly sigmoid, 40-90 n wide and approximately 0-7 mm long ; lacking terminal expansions. Spermathecae 0-6-0-7 mm long ; the duct muscular, approximately one sixth of the length of the ampulla. Ampulla digitiform, wider ectally. DISTRIBUTION. Argentina : Buenos Aires (Barracas do Sul, under stones, on the banks of a river). MATERIAL EXAMINED. 5 semimature syntypes, Buenos Ayres, Beddard collection, B.M. (N.H.) 1904. 10.5. 929/33. Many semimature syntypes, Buenos Ayres, " Barracas d. Sul, Fluss Ufer " collector Michaelsen, 26. VII. 1893, Hamburg Museum, V. 4103. REMARKS. Re-examination of the specimens in the British Museum permits the above very considerable extension of previous accounts. E. rosae is morphologically and probably cladistically very close to E. saltensis but is clearly distinguished by the very short spermathecal ducts and ectal widening of the ampullae. Conspicuous nephropores distinguish the British Museum material from E. saltensis, in which none of the many specimens which have been described possessed visible pores, but nephropores are not recognizable in the many Hamburg Museum specimens. As the specimens in both museums are labelled as types it seems possible that differences in the method and condition of preservation have resulted in the difference in visibility of the pores. Only a brief examination of the Hamburg specimens has been possible, little being observed beyond the form of the spermathecae, wider ectally than entally with an extremely short duct, and the form of the male genital field. Eukerria rubra (Friend, 1916) Kerria rubra Friend, 1916 : 147, Fig. 1-6. 1 = 38 mm, w — 2 mm, s = 90. Setae : dd less than 0-5 u. Clitellum saddle- shaped, XIII-XX. Prostatic papillae inconspicuous, on XVII and XIX. Male pores on XVIII, not in line with the prostate pores. Spermathecal pores 2 pairs, in 7/8 and 8/9, in cd lines (also said to be immediately below c). Septal glands extending between IV and VIII. Gizzard absent. Calciferous glands pear-shaped, arising laterally and disposed ventrolaterally ; apparently with 152 B. G. M. JAMIESON a rather narrow lumen and thick walls. Intestinal origin in XII. Nephridia com- mencing in VII ; absent from segments XI and XIV. Testes and funnels (free?) in X ; seminal vesicles in IX and XI. Vasa deferentia apparently lacking terminal dilatation. Prostates with glandular part lined by a single layer of cells, extending at least to XXI ; ducts short, approximately equal in length to a segment ; lacking terminal bursa. Ovaries and funnels in XIII ; ovisacs in XIV. Spermathecae " pear or bottle-shaped" with slightly swollen ampullae; and slightly longer, fairly sharply demarcated, tubular ducts about half as wide. DISTRIBUTION. Focus of endemicity unknown. Type-locality the Lily House, Oxford Botanical Garden, England, " in oozy mud which surrounded the plants on one side of the tank ". REMARKS. The description of this species, specimens of which are no longer traceable, is inadequate and it probably should be regarded as a species dubium. Eukerria saltensis (Beddard, 1895) Fig. 4I-L, gJ,K, loD Kerria saltensis Beddard, 1895 : 225 ; Beddard, 1896 : 42 ; Michaelsen, 1898 : 479 ; 1900 : 371 ; 1904 : 286 ; 1907 : 23 ; i935a : 103 ; i935b : 40 ; Pickford, 1928 : 378, Fig. 1-4 ; Gates, 1942 : 73 ; Gavrilov, 1952 : 692 ; Jamieson, 1967 : 61, Fig. i. Acanthodnlus sydneyensis Sweet, 1900 : 124, PI. 14, fig. 7, PL 15, fig. 18. Kerria gunningi Michaelsen, 19135 : i, Fig. i ; Michaelsen, 19130 : 419 ; 19136 : 276. 1 Kerria nichollsi Jackson, 1931 : 121, PI. XVI, fig. 5, 8, 9, n. 1 = 25-100 mm, w = 1-2 mm, s = 118-135. Epilobous. In segment XII, aa ca. = be, ab = cd, dd : u = 0-35-0-39 (-0-5?) ; in the type (postclitellar) aa : ab : be : cd : dd = 4 : i-o : 4-4 : i-o : 10-9 ; dd : u = 0-39. Setae a present throughout the clitellum ; setae b present or absent in XVII-XIX ; lateral (and ventral?) setae in the forebody bearing minute teeth. Clitellum annular but less tumescent ventrally, £ XIII, (XIV)-(XIX), (i- XX, XX), | XX (= 7-8 segments). Prostatic pores on minute papillae, on XVII and XIX, considerably lateral of setal lines b of adjacent segments ; those of a side connected by a seminal groove with tumid margins which bends medially in XVIII in which it contains the male pore (i.e. male pores con- siderably mediad to the prostatic pores but still lateral of b lines). Each prostatic papilla encircled by the groove and situated on the summit of a tranvsersely oval prominence which is not clearly denned medially and is in turn borne on a low, earlike prominence which is only laterally elevated. Nephropores not visible externally. Female pores conspicuous or not, anterior in XIV, in b lines, or much less commonly in ab lines or lateral of b lines ; on minute cones or with narrow lips. Spermathecal pores in 7/8 and 8/9, mostly at 2/3 be, occasionally at mid be ; usually readily observed on close examination but never conspicuous. Last septal glands in VI. Gizzard weakly to well developed in VII. Calciferous glands slenderly pear-shaped rather thick walled, permeated by blood vessels (and intracellular spaces?) but without internal folds. Intestine commencing in XII. Hearts 3 pairs ; latero-oesophageal in X and XI ; dorso ventral in IX. Nephridia THE OLIGOCHAETE GENUS EUKERRIA 153 commencing in VI (?), entering the parietes at mid be (with small terminal dilatation of the duct?). Testes and funnels large, free, in X. Seminal vesicles in IX and XI or XI only. Prostates very slender (0-06-0-1 mm wide), winding posteriorly into XXVI or further ; ducts slightly or much more slender, demarcated by their muscular sheen. Neither prostatic nor sperm ducts notably thickened ectally. Ovaries (palmate) and funnels in XIII ; ovisacs absent. Spermathecal ampulla large and oblong-ovoid with thin walls ; duct (always?) with a capacious thin walled ental chamber approximately one third the length of the ampulla, and a muscular terminal portion which is as long as, or shorter than the remainder. Total length of sperm- atheca 0-5-0-8 mm. DISTRIBUTION. South America : Chile ; Juan Fernandez Is ; Argentina. South Africa : Cape Province ; Natal ; Transvaal-Orange Free State border. Burma. New Caledonia. Australia : New South Wales ; S. West Australia (?) ; Queensland. MATERIAL EXAMINED. A single fragmentary previously dissected clitellate type- specimen of K. saltensis, Beddard collection, B.M. (N.H.), 1904, 10.5. 928. A clitellate type-specimen of Kerria gunningi, Hamburg Museum, v. 7490. REMARKS. This is the most widely peregrine species of Eukerria. The type-specimen in the British Museum now yields little information beyond what is indicated in the accompanying illustrations. The prostomium is epilobous \, closed acute ; setal ratios are as recorded above ; nephridia enter the parietes slightly above mid be and have each a very small ectal dilatation of the duct ; the gizzard is about twice the width of the oesophagus but strongly muscular ; the spermathecae have been severed shortly ental to the duct which is muscular and spindle-shaped ; and the prostates are much coiled and extend to XX. I follow Pickford (1928) in including the South African K. gunningi as a junior synonym of E. saltensis. A Hamburg Museum specimen differs from the type of the latter taxon in having a long U-shaped muscular spermathecal duct (the knoblike ampulla being one fourth of its length) but that illustrated by Pickford showed the usual saltensis-iorm and there is no evidence to suggest that the variation observed is not intraspecific. A large number of specimens from South Africa described by Jamieson (1967) accord closely with the type and with Beddard's descriptions of it. Eukerria stagnalis (Kinberg, 1867) Fig. 5, 6, loC Mandane stagnalis Kinberg, 1867 : 100. Acanthodrilus stagnalis ; Vaillant, 1899 : 177. Kerria stagnalis ; Michaelsen, 1899 : 426 ; 1900 : 370. Eukerria stagnalis ; Cordero, 1942 : 278, Fig. 8. Acanthodrilus spegazzinii Rosa, 1890 : 516, i Fig. Kerria spegazzinii ; Rosa, 18955 : 146 ; Beddard, 1896 : 40. 1 = 31-86 mm, w = 0-8-3 mm> s =80-168. Prostomium epilobous $, sometimes £, usually open. Pigmentless in alcohol. Setae closely paired. In segment XII : I54 B. G. M. JAMIESON ("large morp „ small , morph I morph Buncos Aires < „ I small aa ab be cd dd dd : u { 4-0 3'2 I-O i-o 5-6 4-0 0-8 0-9 18-6 15-5 0-50 0-51 3-2 I-O 4-6 I-O 15-6 0-49 { 5-6 I-O 6-8 1-2 16-2 0-41 4-1 I-O 5'5 I-O 17-3 0-48 Paraguay mean of 5 Setae a present, b present or absent in XVIII, totally absent in XVII and XIX ; penial setae absent. Nephropores rarely visible as white dots shortly median of setae c. Clitellum annular though interrupted by the male genital field, occupying XIII \ XIII-XIX, \, f XX (— 6^-7^ segments) ; ventrally less strongly developed and sometimes embayed almost to \ XIV ; intersegmental furrows present only ventrally, setae retained. Male genital field : prostatic pores two pairs of conspicu- ous transverse gaping slits, in XVII and XIX, wider than a setal couple, their centres in line with the ventral setal couples of neighbouring segments ; a single or double conical penis-like structure may be visible projecting through a pore. Each pore almost spanning a low oval papilla which is surrounded by a broad low tumid area which extends laterally to almost mid be, fills the segment longitudinally and is united with those of the other side. Male pores a pair of small, rarely visible slits, on XVIII, shortly lateral of, less commonly at the sites of, setae b ; the ventral couples often translocated medially ; bordered by tumid longitudinal bands, con- fluent or contiguous medially, which are continuous with the tumid prostatic fields. Seminal grooves indistinct tracts connecting the prostatic and male pores of each side or not distinguishable their courses varying according as the male pores are median to, in line with, or lateral to the prostatic pores. Glandular mounds, approximately as large as the prostatic porophores, present in line with or median to the latter postsetally in XVI and presetally in XX, or vestigial or absent, surrounded by tumid areas confluent with the male genital field. Female pores inconspicuous, shortly lateral of b lines anteriorly in XIV. Sperm- athecal pores 2 pairs of transverse slits with slightly raised margins, in 7/8 and 8/9, their centres in or slightly lateral of b lines. Dorsal pores absent. Pharynx in III, invested by lobulated pharyngeal glands which extend to the anterior region of VI. Gizzard totally absent. Calciferous glands ; broadly pyriform, narrowing posteriorly to a short duct which joins the lateral aspect of the oesophagus ; the two glands contiguous or nearly so below the oesophagus. The walls thin, with approximately twelve thick vascularized radial septa projecting into the lumen for varying distances, some uniting with neighbouring septa or with those of the other side or all free ; walls and septa ciliated. Intestine beginning, with abrupt expansion, in XII ; typhlosole absent. Dorsal blood vessel slender in the region between the pharynx and the posterior hearts, not certainly traced onto the pharynx. Dorso ventral commissural vessels in (V?), VI-XI ; those in X-XI forming latero- oesophageal hearts, receiving connectives from the dorsal and supraoesophageal vessels. Supra-oesophageal vessel as wide as or much narrower than the dorsal THE OLIGOCHAETE GENUS EUKERRIA 155 156 B. G. M. JAMIESON vessel, arising anteriorly as a vessel (calciferous vessel) from each calciferous gland, and ending by bifurcation to form the connectives to the hearts of XI ; the cal- ciferous vessels apparently in one specimen giving connectives to the hearts of IX. Calciferous gland on each side supplied apically by a longitudinal latero-oesophageal vessel which is separate from the oesophagus but median to the hearts (traced in VI to IX). Subneural vessel absent. Nephridia stomate holonephridia throughout, the first postseptale in III ; (always?) absent from XIV-XVI ; ducts avesiculate though not especially narrowing. Dense villiform testes, large anterodorsally directed iridescent funnels and sperm masses in X only, enclosed in a delicate circumoesophageal testis-sac which encloses also the hearts and nephridia. Very large dorsally apposed racemose seminal vesicles in XI, attached to its anterior septum ; smaller, much-dissected seminal vesicles in IX attached to its posterior septum. Prostates 2 pairs, the tubular glandular portion slightly de- pressed, tortuous, and extending posteriorly through many segments ; their ducts with a muscular sheen and continuous with the glands by a non-glossy transitional region. Each duct ending ectally at the postero-dorsal aspect of a large muscular hemispheroidal bursa. Widths of the bursae 0-38-0-78 mm (see Remarks). Vasa deferentia united on each side and ectally expanded to form an approximately fusiform or subspherical bursa ; the expansion sometimes extending for much of the length of a segment. Ovaries paddle-shaped laminae composed of linear series of oocytes. Female funnels large and compact ; ovisacs absent. Spermathecae inflated sacs each usually once bent on itself and with a firmer ectally narrowing, usually poorly demarcated duct which is from a quarter to a half as long as the saccular ampulla ; diverticula absent ; total length of a spermatheca of IX 0-9-2-0 mm (see Remarks). DISTRIBUTION. Uruguay : Montevideo (type-locality). Argentina : Buenos Aires. Brazil : Rio Grande do Sul, Porto Alegre. (Limnic). MATERIAL EXAMINED, n clitellate syntypes of K. spegazzini of which i large and i small specimen were dissected ; Buenos Aires, collector L. Borelli, 1893, Torino Museum, ol. 114, ex. 291. 3 clitellate specimens of " Kerria " of which i was dis- sected, N. Paraguay, collector? Hamburg Museum, V. 6713. REMARKS. The existence of infraspecific morphs in E. stagnalis poses interesting problems. It appears unlikely that the large morph from Buenos Aires is capable of pairing with the exactly sympatric small morph and the series of both is sufficiently large to cast doubt on the existence of specimens of intermediate size in the neigh- bourhood. On the other hand the Paraguayan specimens are intermediate in size between the two morphs, their intermediate nature being reminiscent of central populations in a Rassen-Kreis. Additional evidence for the distinctness of the large and small Argentinian morphs is the vestigial nature of accessory markings in the former which are well developed in all 6 clitellate specimens of the small morph and in the 3 clitellate specimens of the Paraguayan morph examined. It is to be hoped that comparisons of ploidy in the three morphs will be undertaken. Comparative data on the three morphs from the small series available are re- corded below. Data are limited as several specimens are posterior amputees or THE OLIGOCHAETE GENUS EUKERRIA 157 gm.b XVII XVII 1*6 FIG. 6. Eukerria stagnalis. A-G, syntypes of Kerria spegazzini, Torino Museum, ol 114, ex. 291 : A-C, large morph, A and B, ventral and lateral aspects of right spermatheca of IX ; C, prostates of same specimen. D-G, small morph, D and E, median and dorsal aspects of left spermatheca of IX ; F, lateral view of calciferous glands ; G, prostates of same specimen ; H-J, " Kerria ", Hamburg Museum, V. 6713, H, prostates ; I and J, ventral and lateral aspects of right spermathecae of VIII and IX. (All to same scale). 158 B. G. M. JAMIESON regenerates and because of the desirability of minimizing dissection. Numbers of specimens examined are shown in parentheses. Measurements are in mm. Small morph Large morph Paraguayan morph Length 31-52 (6) 82-86 (2) 62-82 (3) Greatest width 0-8-1-1(6) 1-5-1-9 (3) 2-5-3(3) Mid-clitellar width 1-8-2-4 3-I-4'4 2-1-2-4 (mean of 4 = 2-1) (mean of 4 = 3-5) (mean of 3 = 2-3) Segments 80-123 (6) 168-170 (2) 88 (regenerating?)— *54 (3) Accessory genital markings Well developed (6) Vestigial (3) Well developed (3) Width prostatic bursae 0-38-0-47(1) 0-69-0-78(1) 0-59-0-69(1) Length of a spermatheca in IX 0-9(1) 2-0(1) 1*2(1) The greater width of the Paraguayan specimens appears to be due to contraction of the forebody, the clitellar width being intermediate between that of the other two morphs. Eukerria subandina (Rosa, 1895) emend. Gavrilov, 1967 Fig. 7, 8, 9H, I, L, loE Kerria subandina Rosa, 18953. : 2 ; Rosa, iSg^b : 143, PI. fig. 17, 18 ; Michaelsen, 1900 : 371 (Including K. borellii) \ Cognetti, 1902 : 3 ; Pickford, 1928 : 381, Fig. 7. Kerria borellii Cognetti, 1900 : 6, PL fig. 6 ; Cognetti, 1902 : 3 ; Pickford, 1928 : 381, Fig. 8 ; Gavrilov, 1967 : 144, Fig. 1-7. 1 = 30-81 mm, w = 1-4-2-4 mm, s = (76?) 100-136, rarely 148-169. Rosy in life. Proepilobous to epilobous. Setae narrowly ornamented from segment II. Setae ab absent or only partly represented in XVII-XIX ; ab = cd (or larger or smaller) ; dd : u = 0-5 or rarely somewhat smaller. In a type of borelli, in XII, aa : ab : be : cd : dd = 3-5 : i-o : 4-4 : 0-63 : 14-3 ; dd : u = 0-47. Nephropores (always?) in be near c ; in the preclitellar segments in front of c ; in the postclitellar segments further presetally. Clitellum annular, less developed ventrally and inter- rupted by the male genital field ; in XII, | XII, * XII-XII, XIII-XIX, £ XX, f XX, | XX, XX (=7-9 segments). Male genital field I-shaped, ventral median between the equators of XVII & XIX ; its angles reaching transversely a variable distance into be ; its median region less extensive, generally not extending above ab. The four prostatic pores central or eccentric on transversely elongated or round papillae, which extend in the angles of the male field, from b laterally. Seminal grooves, with more or less elevated and whitish margins, convex towards the mid- ventral line, reaching to or approximately to a ; male pore at the bottom of each groove, at the equator of XVIII with a tendency to be very slightly lateral of a. Prostatic papillae and the areas delimiting them, joining the elevations of the seminal grooves, constituting little-developed porophores which laterally delimit the male genital field ; the area median to these, with anterior and posterior margins of variable configuration, may be prominent, level with the remainder of the integument, or depressed. In several re-examined syntypes of borelli there is a pair of pad-like accessory genital markings at 19/20, sometimes represented on one side only or THE OLIGOCHAETE GENUS EUKERRIA 159 pr. po1 XVII XVIII — - XIX— pr. po2 pr. po1 pr.g. FIG. 7. Eukerria subandina. A-E, syntypes of Kerria borelli, Torino Museum, ol. 115, ex. 467 : A, right spermatheca of VIII ; B, lateral and C, ventral views of the male genital field of the same specimen ; D, previously excised male genital field of another specimen ; E, prostates of first specimen. F and G, syntype of K. borelli, Hamburg Museum, V. 5896 : male genital field. 160 B. G. M. JAMIESON absent. Female pores at the anterior margin of XIV, a certain distance behind 13/14, in or lateral to b lines. Spermathecal pores inconspicuous or on papillae, in 7/8 and 8/9, at mid be or as far dorsally as c lines. Last septal glands in VI or VII. Gizzard relatively well developed, in VII ; its muscular coat 3-5 times as thick as the internal epithelium. Calciferous glands (re-examination of the type of borelli) with thick walls traversed by longitudinal blood sinuses, which occupy the entire width of the walls, between which are narrow canaliculi which probably are continuous with the central lumen ; septa absent; structure similar in a type of subandina. Intestine beginning in XII, generally, at ii 1 12 ; typhlosole absent. Dorsal and ventral vessels single ; supraoesophageal from anterior XI to posterior IX ; subneural and extraneurals absent. Hearts in IX-XI ; lateral in IX ; laterooesophageal in X & XI ; dorsoventral commissurals seen in VIII, in one syntype of borelli, but absent from VII. Nephridia from V (occurring at least as far forward as VI in a re-examined syntype), peritoneum highly developed from the beginning of the clitellum. Testes free ; seminal vesicles race- mose, 2 pairs, in IX & XI. Prostates 2 pairs, tubular, convoluted and folded, ex- tending backwards to XXI-XXXVII. Ducts much thinner than and well differ- entiated from the glandular parts, relatively short, equivalent in length to 1^-2 segments. Ovaries and funnels in XIII ; ovisacs (sometimes rudimentary or absent) in XIV. Spermathecae adiverticulate, claviform, digitiform or inverted pyriform, with a thick, relatively short duct (j-^ of the length of the ampulla) ; length of the right spermatheca of IX in a re-examined syntype = i-i mm ; of the right spermatheca of VIII in a re-examined syntype of borelli = i-o mm. DISTRIBUTION. Argentina. Bolivia. Brazil. MATERIAL EXAMINED. 5 clitellate syntypes of K. subandina (i lacking the preclitellar portion) of which i was dissected, Salta, Argentina, collector L. Borelli, 1893 ; Torino Museum, ol. 115, ex. 467. 8 clitellate syntypes of Kerria borelli of which i was dissected, Urucum, Matto Grosso, Brazil, collector L. Borelli, 1899 ; Torino Museum, ol. 106, ex. 497. i clitellate syntype of K. borelli (same locality data) ; Hamburg Museum, v. 5896 (not dissected). REMARKS. After a careful examination of new material from Arroyo del Toro, Province of Tucuman, Argentina, Gavrilov (1967) accepted the suggestion of Michaelsen (1900) that E. borelli is a synonym of E. subandina. He demonstrated uniparental reproduction in this species. The writer's examinations of type- material of both taxa have yielded no evidence which invalidates regarding borelli as a junior synonym and variation falls in every detail within that indicated in the synoptic description of Gavrilov (1967) summarized above. Accessory genital markings noted unilaterally or on both sides in 19/20 in five of six Torino syntypes were not, however, described by Gavrilov. The markings were absent, or perhaps represented by a medianly continuous slight rim or pad, in one of the six specimens and in the Hamburg Museum specimen. The albeit inconsistent occurrence of these markings and the conspicuous appearance of the spermathecal pores confirmed in the re-examination both contrast with the condition in Argentinian specimens and the possibility of subspecific distinction of Brazilian populations deserves investigation. THE OLIGOCHAETE GENUS EUKERRIA 161 _ in 2 o| — • £ nj C of " 3 t^,Q vO fl *a x' ffi M 0 .> « -t-> r< O 1 21 QX I I-H ^tj SH „ O °l s i ^ U o £ 162 B. G. M. JAMIESON Eukerria tucumana Cordero, 1942 Fig. QM, N Eukerria tucumana Cordero, 1942 : 281, Fig. 12-15. 1 = 63-65 mm, w = 1-5-2 mm, s = 105-120. Epilobous. Setae : ab = cd ; aa less than be anteriorly, = be posteriorly ; aa = 4-5-5 ab and dd>o-$ u, especially in the anterior portion. Nephropores in be, nearer c. Clitellum saddle-shaped, XIII-XIX, \ XX. Male genital field between XV and XX which region is de- pressed and concave. 2 pairs of oval tubercles in the most depressed region (XVII- XX), traversed longitudinally by seminal grooves the borders of which are strongly tumid, especially laterally ; prostatic pores in the setal arcs of XVII and XIX slightly lateral of setae b, at the ends of the seminal grooves, on simple papillae which do not project markedly ; the seminal grooves linking these with the male pores, which are in the same longitudinal line at the middle of XVIII which is much extended and depressed. Each pair of tubercles crossed transversely by the inter- segmental furrows bordering segment XVIII so that one third lies in XVIII. Female pores anterior in XIV, anteromedial to setae b. Spermathecal pores in 7/8 and 8/9, in b lines, recognizable by a slight increase in the parietal pigmentation in their vicinity. Last septal glands in VI. Gizzard well developed ; at i mm wide, much wider than the oesophagus. Calciferous glands arising dorsally from the oesophagus ; with long curved ducts ; the sacs below the gut ; as in Ocnerodrilus with small parietal septa. Intestine commencing in XIII. Testes and funnels in X, (free?) ; seminal vesicles racemose, in IX and XI. Prostates extending to XXIX, very attenuated ; their ducts more slender, a little more than the length of a segment ; without terminal expansions. Ovaries in XIII. Spermathecal ampulla triangular, its apex con- tinuing without constriction as a long, wide, coiled duct. DISTRIBUTION. Argentina : Tucuman. (Habitat?). REMARKS. E. tucumana shows affinities with E. pascuorum and E. eiseniana. Eukerria urna Righi, 1968 Fig. 90 Eukerria urna Righi, 1968 : 183, Fig. 6-8. 1 = 227-28-4 mm, w = 1-23-1-41 mm, s — 76-89. Zygolobous. Reddish pink in life. Setae sigmoid with small, irregularly arranged longitudinal furrows ; aa : ab :bc : cd : dd = 5-3 : i-o : 6-7 : i-o : 14-3, in the midbody, = 3-1 : i-o : 4-2 : i-i : 8-6, in the hindbody ; dd : u — 0-39 and 0-35 respectively (computed as 0-5 by Righi). Nephropores not visible. Clitellum annular, less developed ventrally, \ XIII, \ XIV-^ XX. Prostatic pores small transverse slits on pointed, mamillate elevations on XVII and XIX in setal lines b or slightly above these ; those of a side connected by a thin-walled, whitish seminal groove ; each groove slightly bent laterally at the male pores, in XVIII. Female pores in the anterior half of XIV, in THE OLIGOCHAETE GENUS EUKERRIA 163 front of setae b. Spermathecal pores mostly unrecognizable, sometimes surrounded by distinct oval fields, in 7/8 and 8/9, sometimes immediately below setal lines b, sometimes in the upper half of be ; in 5% of worms (136 clitellate specimens exam- ined) the pores in 7/8 are in the upper half of be and those of 8/9 shortly below c lines. Gizzard strong, clearly distinguished from the oesophagus. Calciferous glands arising laterally from the oesophagus, in IX, rounded, of the ocnerodriloid type. Intestine commencing in XII. Last hearts in XI, lateral. Testes, sperm masses and funnels free in X ; seminal vesicles in IX and XI, the latter pair displacing 11/12 and sometimes 12/13 posteriorly. Prostates with an irregular course below the gut, ending between XXIII and XXVIII ; duct much thinner than the glandular part, of variable length, penetrating only a single septum or extending through 3 segments, i pair of ovaries and funnels, in XIII. Spermathecal duct mostly thinner than the broadly oval ampulla and somewhat longer, bent in various ways. DISTRIBUTION. Brazil : Marajo Island, at Cachoeira do Arari (banks of a river). REMARKS. Clearly this species is close to E. saltensis, as Righi has stated, al- though the form of the spermathecae and the location of the male pores lateral, rather than median of, the prostatic pores clearly distinguish it from the latter species and the internal structure of the calciferous glands is apparently distinct. Eukerria weyenberghi Cordero, 1942 Eukerria weyenberghi Cordero, 1942 : 279, Fig. 9-11. 1 = 37-68 mm, w = 3-4 mm, s = 59-109. Epilobous. Setae : ab = cd, aa = be = 3 ab ; dd : u — 0-5. Nephropores in c lines. Clitellum annular, XIII, \ XIII-XX ; some or nearly all of the ventral and dorsal couples may be obscured ; all intersegmental furrows obscured except 13/14 which is partially visible. Prostatic pores two pairs of widely open elliptical slits level with the general body surface, not on papillae, surrounded simply by an " eyelid " like zone formed solely by modifica- tion of the cuticle ; the centres of the pores in b lines, their internal margins in a. Seminal grooves absent. Male pores visible with difficulty on XVIII in line with the external margins of the prostatic pores and midway between the latter. Female pores on XIV near the anterior border, in b lines at the bottom of a little marked transverse furrow. Spermathecal pores visible on separation of the borders of the intersegmental furrows 7/8 and 8/9 as minute simple orifices in b lines. Last septal glands in VII, in which the gizzard is present though no thicker than the remainder of the oesophagus. Calciferous glands " grape-seed shaped ", arising ventrally from the oesophagus and extending anteriorly ; internally with a central cavity with radial septa inserted on its periphery. Last hearts in XL Testes and funnels free in X ; seminal vesicles one pair, in IX. Prostates longer than those of any other known species, extending in situ to XXXII, the sinuous glandular region, which is quadrangular in section, occupying a length of 8 mm (its actual length about 3 times this) ; duct somewhat narrower, circular in section, smooth and muscular and coiled in a spiral, extending into XXII ; about one eighth the length of the glandular portion. Prostatic ducts discharging on muscular hemispherical papillae 164 B. G. M. JAMIESON G prpl pr.p2 L M N O FIG. 9. Eukerria, reviewed illustrations of male genital fields. A, E. halophila (after Beddard, 1892) ; B-D, E. kukenthali : B and C, E. asilis (after Righi, 1968) ; D, syntype (after Michaelsen, 1908) ; E and F, E. mcdonaldi (after Eisen, 1893) ; G, E. papillifera (after Rosa, 18950) ; H, E. subandina (= Kerria borelli) (after Cognetti, 1900) ; I, E. subandina (after Gavrilov, 1967) ; J, K, E. saltensis (after Jamieson, 1967) ; L. E. suban- dina (after Rosa, 18950) ; M and N, E. tucumana (after Cordero, 1942) ; E. urna (after Righi, 1968). THE OLIGOCHAETE GENUS EUKERRIA 165 corresponding with the external pores and containing a large " atrial " chamber. Prostatic duct joining the postero-medial aspect of the internal papilla. Sperm- athecae 3 mm long by i mm wide ; the ampulla large and pyriform ; each flexing around the oesophagus which is in contact with the duct. The duct wide and ampulliform, distinguishable by its texture and greater opacity ; equalling the ampulla in length. DISTRIBUTION. Argentina : Buenos Aires Province, Islas del Tigre (Habitat?). REMARKS. It seems probable that the internal papillae (bursae?) at the ends of the prostatic ducts are capable of eversion or protrusion to give external papillae of the type seen in other species of Eukerria. DISCUSSION Within the tribe Ocnerodrilini (= Ocnerodrilinae s. Gates, 1966), only Eukerria, Kerriona Michaelsen, 1924, and Maheina Michaelsen, iSggb, display the acantho- drilin condition of the male pores, with the prostatic pores on XVII and XIX and the openings of the vasa deferentia intermediate on XVIII. An especially close re- lationship between Maheina and Eukerria can be rejected as the single species of Maheina differs from Eukerria in its setal ratios (the setae of the fore- and mid-body being widely separated), in location of the gizzard in VI, in possessing two pairs of calciferous glands ; in having testes in X and XI, and geographically, being the only Ocnerodriline known from the Seychelles. A close relationship between Eukerria and the two known species of the Brazilian genus Kerriona was proposed by Stephenson (1930) as Kerriona besides having acanthodrilin male terminalia has the testes confined to segment X as in Eukerria, such proandry being known elsewhere in the Ocnerodrilinae (s. lat) only in Haplo- drilus Eisen, 1900 and, now, in Gatesia Jamieson, 1962. Contrary to the views of Michaelsen (1924) and Stephenson (1930), the terrestrial mode of life of Kerriona cannot be considered a valid distinction from Eukerria as some species of the latter genus are known only from terrestrial habitats. Nevertheless, Kerriona shows morphological distinctions which set it apart from Eukerria and which suggest that the mutual possession of proandry and acanthodrilin male terminalia in the two genera does not indicate a closer relationship between the two genera than either has with other genera of the Ocnerodrilini. Of the few known characteristics of Kerriona, those which indicate that it is phyletically and phenetically distinct from Eukerria are wide pairing of setae in at least the mid and hindbody ; the panicled- tubular or tubular calciferous glands, and the presence of an intestinal typhlosole. In the subfamily Octochaetinae variation from the acanthodrilin condition to the microscolecin condition (a single pair of male and prostatic pores, on XVII) of many ocnerodriles occurs within a single genus, Lennogaster, and therefore the possibility of close relationship of Eukerria and non-acanthodrilin ocnerodriles deserves attention. At present there is, however, no convincing evidence for such a relation- ship though Gates (1957), in a key to the genera of the Ocnerodrilinae, placed those species of Eukerria which lack gizzards in an Ocnerodrilus-group of species. He stated that Eukerria must be restricted to those species with a gizzard in segment VII 166 B. G. M. JAMIESON and went so far as to specify that " Kerriona may be closer to Ocnerodrilus than to its supposed ancester Kerria ". With regard to relationships within Eukerria there can be no a priori justification for segregation of species which lack a gizzard from the remainder of a genus in which development of the gizzard varies from weak to strong, as Jamieson (1963) showed for the genus Nannodrilus, but perusal of the accounts given above in the systematics section does reveal that absence of gizzards correlates with other distinctions in one group of species. Some grounds therefore exist for recognizing subgroups within Eukerria though elevation of these to generic rank seems inadvisable in view of the many gaps in our knowledge of the genus. This agiceriate species-group is the only clearly defined subgroup which the author is able to recognize from the present limited evidence. It may be termed the stagnalis-group and contains only E. stagnalis, E. papillifera and E. weyenberghi. These species share a number of characters which are individually or at least in com- bination very distinctive. They are : (i) absence of a gizzard, at least as a recognizable swelling of the oesophagus (only in E. rubra, elsewhere in the genus, is a gizzard said to be absent) ; (ii) location of the prostatic pores in line with the ventral setal couples (a con- dition occurring elsewhere in E. halophila, E. kukenthali, E. tucumana and E. urna) ; (iii) presence on the prostatic ducts of ectal bursae, which do not occur elsewhere in the genus ; (iv) the absence (E. papillifera and E. weyenberghi} or slight development (E. stagnalis) of seminal grooves (doubtfully absent in E. halophila and E. rubra) ; (v) extension of the prostatic glands through many segments, a feature seen also in E. kukenthali. The structure of the calciferous glands is not known for papillifera but in weyen- berghi small parietal septa were seen by Cordero (1942), a condition which does not conflict with that in E. stagnalis (p. 154) in which, however, some septa are known to fuse centrally. Cordero's statement that in weyenberghi a gizzard is present in VII, though no wider than the oesophagus, would, if correct, suggest real affinity of the stagnalis group with other species of Eukerria. The extensive series of precardiac commissural vessels in E. stagnalis appears to be a primitive feature but, from the evidence of weyenberghi, it appears more likely that absence of the gizzard in stagnalis and papillifera is secondary. Subgeneric or even separate generic status for the stagnalis group is not without justification but the apparently close relationship with Eukerria and the paucity of our knowledge of the latter do not warrant making such distinctions at present. In the absence of taxonomically important information with regard to several systems in many species of Eukerria, the character which appears most likely to permit subdivision of the remaining Eukerrias into morphologically and, presumably, phylogenetically distinct groups is the internal structure of the calciferous glands. Three categories may be distinguished on the basis of this character as shown below. THE OLIGOCHAETE GENUS EUKERRIA 167 STRUCTURE OF THE CALCIFEROUS GLANDS IN EUKERRIA * personal examination (I) Walls very thick and not projecting as septa or as folds into the lumen (Fig. loE) E. limosa E. pascuorum E. subandina* and its junior synonym E. borelli* E. rubra? (II) Transitional. Walls very thick but with a few longitudinal folds though with no denned complete or incomplete septa (Fig. loD) E. rosae E. saltensis* (III) Walls relatively thin. Parietal septa well denned and numerous (Fig. loA-C) E. eiseniana* and its junior synonym E, hortensis E. garmani* E. halophila? E. kukenthali E. tucumana E. urna E. mcdonaldi? (or II) and the stagnaUs— group species, E. stagnalis*, E. weyenberghi. Internal structure unknown. E. asuncionis and E. papillifera. Information on several of the species listed is inadequate and the extent of in- dividual variation needs to be investigated but this albeit crude classification serves to suggest a starting point for further subdivision of the genus. After removal of the stagnalis-group it is not possible to place all the remaining species in subgroups though some groupings are observable. In group I, E. limosa and E. pascuorum are morphologically similar or perhaps synonymous. The two species of group II, E. saltensis and E. rosae appear to be closely related though distinct species. Of the group III species, few affinities are discernible, again largely because of lack of data. E. eiseniana and E. garmani are mutually close but the glands of eiseniana are larger and have more delicate and more numerous septa than those of garmani. E. mcdonaldi isolated in Baja California, stands apart in possess- ing a type of spermathecal diverticulum. Affinities of the other species, E. tucumana, E. urna and the type-species, E. halophila with other species are uncertain though they must at least for the time being be regarded as congeneric. E. urna resembles E. saltensis more than it does other species but its ocnerodriloid calciferous glands are a noteworthy difference. E. tucumana appears to have its closest affinities with E. garmani. It is hoped that drawing together our limited knowledge of Eukerria in this account will stimulate further investigation of the genus, ideally by workers in South America, and that sufficient data will be forthcoming to permit a taxonometric investigation of the affinities of its species. The species of Eukerria recognized as valid in the present work, their junior i68 B. G. M. JAMIESON sep FIG. 10. Calciferous glands in transverse section. A, Eukerria eiseniana, syntype, Torino Museum, ol. no, ex. 295 ; B, E. garmani (garmani), syntype, Torino Museum, ol. in ; C, E. stagnalis, syntype (small morph) of E. spegazzini, Torino Museum, ol. 114, ex. 191, showing two glands and the oesophagus ; D, E. saltensis, specimen from the Great Berg River, South Africa, author's collections ; E, E. subandina, syntype of E. borelli, Torino Museum, ol. 106, ex. 497. (All to the same scale). THE OLIGOCHAETE GENUS EUKERRIA 169 synonyms, their distribution, and the sources of material examined are set out in the taxonomic summary below. TAXONOMIC SUMMARY Species here recognized 1. E. asuncionis (Rosa, 1895) 2. E. eiseniana (Rosa, 1895) Junior Synonyms + = new synonymy 3. E. garmani (Rosa, 1895) 4. E. halophila (Beddard, 1892) 5. E. kukenthali (Michaelsen, 1908) K. hortensis Stephenson, 193 1* K. selangorensis Stephenson, 1931 E. peguana Gates, 1 942 + E. asilis Righi, 1968 + 6. E. litnosa (Stephenson, 1931) 7. E. mcdonaldi (Eisen, 1893) K. zonalis Eisen, 1893 + 8. E. papillifera (Rosa, 1895) 9. E. pascuorum (Stephenson, 1931) 10. E. rosae (Beddard, 1895) U.S. rubra (Friend, 1916) 12. E. saltensis (Beddard, 1895) Museum material examined B.M. = British Museum T. = Torino Museum H. = Hamburg Museum * = Type material T* T* B.M.* B.M. T* Distribution B.M. H* B.M.* B.M.* T* B.M.* B.M.* T* H* B.M.* A canthodrilus Sydney ensis — Sweet, 1900 K. gunningi H* Michaelsen, 1913 K. nichollsi — Jackson, 1931 Paraguay Paraguay Paraguay Paraguay; Argentina Bolivia(?) : upper reaches of the Pilcomayo St. Thomas, West Indies ; Christmas Island, Indian Ocean Selangor, Malaya Burma Brazil : Marajo Island Paraguay Baja California Baja California Paraguay Paraguay Argentina Oxford botanical gardens Chile-mainland and Juan Fernandez Is ; Argen- tina ; Burma ; S. Africa ; Queensland New South Wales ; Vic? South Africa ; New Cale- donia South West Australia iyo B. G. M. JAMIESON 13. E. stagnalis (Kinberg, 1867) 14. E. subandina (Rosa, 1895) 15. E. tucumana Cordero, 1942 1 6. E. urna Righi, 1968 17. E. weyenberghi Cordero, 1942 TAXONOMIC SUMMARY (continued) Acanthodrilus spegazzini T* H. Rosa, 1890 K. borelli Cognetti, 1900 T* T* H* Uruguay Argentina ; Brazil Argentina Brazil Argentina Brazil : Marajo Island Argentina ACKNOWLEDGEMENTS This study was made possible through the kind cooperation of Mr. R. W. Sims, British Museum (Natural History), Dr. M. Dzwillo, Zoologisches Museum, Hamburg, and Dr. L. Parenti of the Museo ed Istituto di Zoologia Systematica, University of Torino to whom the author gratefully extends his thanks. Serial sections were prepared by Mr. J. Casey. The work was financed by the Canadian National Research Council and University of Queensland Research Grants. Special thanks are due to Professor R. O. Brinkhurst for facilities provided. ILLUSTRATIONS With the exception of Fig. 9, the illustrations have been drawn by the author by camera lucida. The scale indicated is I mm unless otherwise labelled. Shaded areas represent the clitellum. Abbreviations used in the illustrations : b.s, blood sinus ; c, cerebral ganglia ; ca. g, calciferous gland ; cil, cilia ; d.v. dorsal blood vessel ; dv. h, dorso ventral (lateral) heart ; ep, epithelium ; $, female pore ; gi. gizzard ; g.m, accessory genital marking ; g.m.b, bursa corresponding with external genital marking ; int. v, intestinal (oesophageal) valve ; lac, cavity or lacuna ; lo, latero-oesophageal vessel lo.h, latero-oesophageal hearts ; <$ male pore ; $ f, seminal funnel ; n, nucleus ; n.c, ventral nerve cord ; np, nephropore ; oe, oesophagus ; ph, pharynx ; pr. d, prostate duct ; pr. g, glandular part of prostate ; pr. b, prostatic bursa ; pr. p, prostate pore ; pr. po, prostate porophore ; pro, prostomium ; sec. gr, secretory granules ; sem. gr, seminal groove ; sep, septum ; sep. g. septal gland ; sp, spermatheca ; sp. p, spermathecal pore ; s.v, seminal vesicle ; t, testis ; v.d, vas deferens ; v.d.b, ectal thickening (bursa) of vas deferens. 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Some new species of Ocnerodrilinae (Oligochaeta). Proc. zool. Soc. Lond., 139 : 607-626. KINBERG, J. G. H. 1866. Annulata nova. Ofvers. K. VetenskAcad. Fork., 23, 4 : 97-103. MICHAELSEN, W. 1898. Die Oligochaeten der Sammlung Plate. Zool. Jb. Supplt., 4 : 471-480. iSgga. Revision der Kenberg'schen Oligochaetentypen. Ofvers. K. VetenskAkad. Fork., 56 : 413-448. — iSggb. Oligochaten von den Inseln des Pacific, nebst Erorterungen zur Systematik der Megascoleciden. Zool. Jb. Syst., 12 : 211-246. — 1900. Das Tierreich, 10, Vermes, Oligochaeta. 1904. Catalogo de los Oligoquetos del territorio chileno-magallanico i descripcion de especies nuevas. Revta chil. Hist, nat., 8 : 262-322. 1907. Oligochaeten von Australien. Abh. Geb. Naturw., Hamburg, 19, i : 1-25. 1908. Die Oligochaten Westindiens. Zool. Jb. Supplt., 11 : 13-32. — igi3a. The Oligochaeta of Natal and Zululand. Ann. Natal Mus., 2, 4 : 397-458. 191 3b. Oligochaten von tropischen und siidlich-subtropischen Afrika. Zoologica, Stuttg., 27, 68 : 1-63. I9I3C. Die Oligochaten von Neu-Caledonien und den benachbarten Inselgruppen. Sarasin, Fritz and Jean Roux, Nova Caledonia. A. Zoologie, 1, 3, 5 : 171-280. 1924. Oligochaten von den warmeren Gebieten Amerikas und des Atlantischen Ozeans sowie ihre faunistischen Beziehungen. Mitt, naturh. Mus. Hamburg, 41 : 71-83. i935a. Oligochaeta from Christmas Island, south of Java. Ann. Mag. nat. Hist. (10), 15, 85 : 100-108. I935b. Earthworms from South Western Australia. /. Proc. R. Soc. West. Aust., 21 : 39-43- PICKFORD, G. E. 1928. Synonymy in the genus Kerria (Oligochaeta, Ocnerodrilinae). Ann. Mag. nat. Hist. (10), 2 : 378-382. RIGHI, G. 1968. t)ber die Oligochatengattung Eukerria. Beitr. neotrop. Fauna, 5 : 178-185. 172 B. G. M. JAMIESON ROSA, D. 1890. Terricoli Argentini raccolti dal Dott. Carlo Spegazzini. Annali Mus. civ. Stor. nat. Giacomo Doria, 29 : 509-521. iSgsa. Oligocheti terricoli. Boll. Musei Zool. Anat. comp. R. Univ. Torino, 10, 204 : 1-3. 18950. Contribute allo studio dei terricoli neotropicali. Memorie Accad. Sci. Torino (2), 45 : 89-152. STEPHENSON, J. 1930. The Oligochaeta. Oxford. 1931. The Oligochaeta from Brazil and Paraguay. /. Linn. Soc., Zool., 37, 252 : 291-326. SWEET, G. 1900. On the structure of the spermiducal glands and associated parts in Austra- lian earthworms. /. Linn. Soc., Zool., 28, 180 : 109-139. VAILLANT, M. L. 1889. Lombriciniens, Hirudiniens, Bdellomorphes, T6retulariens et Planariens. Quatrefages de Bre"au, J.L.A. de. Histoire naturelle des Annelds marins et d'eau douce, 3, i : I-XII : 1-340. BARRIE G. M. JAMIESON, Ph.D. Zoology Department UNIVERSITY OF QUEENSLAND ST. LUCIA BRISBANE QUEENSLAND, AUSTRALIA Printed in Great Britain by Alden & Mowbray Ltd at the Alden Press, Oxford OBSERVATIONS ON THE ELECTRA DOLPHIN, PEPONOCEPHALA ELECTRA W. A. DAWBIN, B. A. NOBLE, F. C. FRASER BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 20 No. 6 LONDON : 1970 OBSERVATIONS ON THE ELECTRA DOLPHIN, PEPONOCEPHALA ELECTRA BY WILLIAM HENRY DAWBIN BRUCE ALEXANDER NOBLE FRANCIS CHARLES FRASER Pp. 173-201 ; 13 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 20 No. 6 LONDON : 1970 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, is issued in five series corresponding to the Departments of the Museum, and an Historical series. Parts will appear at irregular intervals as they become ready. Volumes will contain about three to four hundred pages, and will not necessarily be completed within one calendar year. In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department. This paper is Vol. 20, No. 6 of the Zoological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. Trustees of the British Museum (Natural History), 1970 World List abbreviation Bull. Br. Mus. nat. Hist. (Zool.). TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 21 December, 1970 Price gop OBSERVATIONS ON THE ELECTRA DOLPHIN, PEPONOCEPHALA ELECTRA By W. H. DAWBIN, B. A. NOBLE & F. C. FRASER INTRODUCTION THE notes set down in this paper were stimulated by a stranding of electra dolphins at Crowdy Heads, about 200 miles north of Sydney in August 1958. This stranding was reported briefly as a school of blackfish by a local newspaper some days after the event. Following notice of the report one of the authors (W.H.D.) made local enquiry and was told that all the animals had been removed by fishermen and had been cut up for bait. The local postmistress, Mrs. W. J. Ward, had made a careful count of the teeth of one specimen before its removal and stated that there were 22 on each side of the upper jaw and 19 on each side of the lower jaw. This eliminated the possibility that the school might have been pilot whales (Globicephala sp.) the species usually referred to locally as blackfish. The observed tooth row number also ruled out Feresa from consideration. An immediate search in the Crowdy Heads area failed to unearth any skeletal remains, so with the co-operation of the Australian Broadcasting Commission, both radio and television, appeals were made for any photographs or material which may have been taken by visitors to the beach on the day of the stranding. The response in colour and black and white photographs and in descriptions was good, and a number of the photographs showed quite clearly that most of the animals had a rounded snout without beak, white lips and light thoracic and abdominal patches on an otherwise uniformly dark-coloured background. During a further visit to the Crowdy Heads area 1962 (by W.H.D.) reports inland from the area where received that in the course of removal of the dolphin carcases one had fallen off a truck and had been left behind at the roadside. The skull had been removed by a local resident, Mr. H. Anderson, who generously presented it to the visiting author. Subsequently a half skull which had been used as bait in a fish trap was secured. This specimen was from a dolphin that had stranded in the Port Macquarie area in January 1962. The unprompted description of the dolphin by the fisherman con- cerned matched those at Crowdy Heads. He had seen the latter animals and was convinced that his specimen was identical with them. The number of alveoli in the skull of the decomposed carcase and in the half skull essentially confirmed Mrs. Ward's tooth count and from this and from other skull characters the school was provisionally identified as Lagenorhynchus (— Pepono- cephala) electra (by W.H.D.). The two specimens were sent to the British Museum (Natural History) for comparison with other examples of this species including the holotype. 176 W. H. DAWBIN, B. A. NOBLE, F. C. FRASER OBSERVATIONS ON THE CROWDY HEADS AND PORT MACQUARIE STRANDINGS (W.H.D.) Letters referring to the animals prior to stranding were unanimous in describing them as a large school of black dolphins sighted off Port Macquarie heading south between 2 and 4 p.m. on the day preceding the stranding. Crowdy Heads is approx- imately 40 miles south of Port Macquarie and the animals reached there and com- menced stranding during a high tide about 10 a.m. the following morning, along a gently shelving stretch of sandy beach. They stranded in more or less a line with heads facing away from the sea and the animals were spread between a quarter and half a mile of beach. Number estimates range from 150 to 250 in the school and none was seen which did not come ashore. A few which were pushed out to sea returned fairly quickly and became stranded again. Length estimates state that the school contained small young about 3 ft in length but were mostly about 8 ft with some possibly up to 10 ft ; however actual measurements do not appear to have been made. A considerable proportion (some writers state up to 50% :) were pregnant females and there was agreement that the foetuses were of similar size estimated as 2'6" to 3'o" and judged by appearance as near full term. A photograph of one foetus held up by a fisherman is consistent with the size estimate. The observations on foetuses and also of apparently new born young suggest a relatively circumscribed breeding period during spring. No observations on stomach contents were reported but there were several reports of small thread-like worms " as a crawling mass in the heads. " These appear to have been common and resemble the report of Nakajima and Nishiwaki (1965) on the presence of many nematodes in the air sinuses of the skull. The occurrence of these animals in a single school of 150-250 and the report of another and larger school off Japan in 1965 (Nishiwaki and Norris, 1966) indicate that this is a schooling species, despite the isolated nature of each of the skulls received. A smaller group (estimated as about two dozen) has been reported by a fisherman (Mr. L. Elford) who collected specimens from the Crowdy Heads school and also obtained the Port Macquarie specimen. These were stranded at Diamond Head, a few miles from Port Macquarie and all were cut up and completely used for bait. Other local fishermen recognised that the size, colour pattern and head shape differed from the usual local dolphins and agreed with those at Crowdy Heads. They also report occasional sightings of similar schools at about the edge of the continental shelf along the neighbouring coast. It is therefore possible that P. electro, is not particularly rare along the coast of northern New South Wales but that it is a species which normally remains some distance from land. External features The earliest representation of the external appearance of the electra dolphin is provided by Peale (1848) and is reproduced in True (1889). The lateral view of the animal shows it to be darkly pigmented over most of the body, but showing a white patch midway between lower jaw and flipper and an elongated white ventral area extending from some way in front of the level of the ELECTRA DOLPHIN 177 anterior margin of the dorsal fin, to a level midway between dorsal fin and tail. There is indication of a white lower lip and lighter pigment around the eye. The next figure of external appearance is that of Owen (1866), (his plates 5, fig. I and 7, figs. 1-5) with reference to L.fusiformis (a synonym). This reference is given only to emphasise the error in representation of the electra dolphin. Goodwin (1945) reproduced the drawing made by Robert Cushman Murphy of an electra dolphin taken in the tropical Atlantic in 1912. This drawing shows an absence of pigmentation around the mouth and on the throat, but no white area more posteriorly on the belly. A noteworthy feature is a ventrally projecting, long, low eminence in the post-anal region. The photographs included in Nishiwaki and Norris (1965) best show the external form and pigmentation. These authors give a description of shape, particularly of the head, in connection with the erection of the genus Peponocephala to include P. electra, because of the absence of a well defined rostrum. The various pictures show the electra to be predominantly darkly pigmented. The white mouth area is conspicuous and in the ventral view the broad, unpigmented throat area behind the deeply pigmented lower jaw area, links up with the white patch seen in lateral view in Peale's and Goodwin's figures. At about the level of the flipper, the dark pig- mentation extends from the side down to the mid ventral line. Behind this, a lenticular unpigmented area, extends from umbilicus to anus ; this area again is the same area seen in restricted lateral view in Peale's figure. Specimens The following specimens were available for survey : — No. Reg. No. Comments 1 B.M. (N.H.) 1844.10.5.3 Type of Lagenorhynchus electra Gray, 1846 Zool. Erebus & Terror, I (Mammalia) 35, pi. 13. [ADULT in growth comparison] 2 B.M. (N.H.)i866.2.5.i Type of Delphinus fusiformis. Owen, 1866 i475a Trans. Zool. Soc. London, 6 : 22, pi. 5, fig. i, pi. 7, figs. 1-5. 3 B.M. (N.H.) 358a Type of L. asia Gray, 1846 Zool. Erebus & Terror I (Mammalia) 35, pi. 14. 4 W.A.M. 4798 Salt marsh at Derby ; W. Australia. Skull and skeleton. 5 A.M.N.H. 4300 c? found at 3°03'N 24°4o'N in Atlantic. Goodwin 1945 J. Mammal 26 : 195. 6 B.M. (N.H.) 1959.7.9.2. c? Hittadu Is., Addu Atoll. Can Maldives. Skull and skeleton. [ADOLESCENT in growth comparison] 7 New South Wales. No lower jaw. [JUVENILE in growth comparison] 8 New South Wales. Half skull, no lower jaw. 9 B.M. 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H. DAWBIN, B. A. NOBLE, F. C. FRASER The post-temporal fossa (fig. 2) is bounded anteriorly by the postorbital process which is contained as a ridge of the exoccipital ; posteriorly the fossa is bounded by a ridge between parietal and exoccipital. TABLE 2 i. 2. 3. 4. 5. 6. 7. 8. 9. 10. n. 12. 13. 14. 15. 1 6. 17- 1 8. 19- 20. 21. 22. 23- 24. 25- 26. 27. 28. 29. 30. 31. 32. 33. 34. 35. Condylo-basal length. Rostrum length. ,, basal width. ,, width 60 mm. anterior to ant. ob. notches. ,, ,, at middle. Premaxillae width at same point Tip of snout to blowhole. ,, ,, ,, ,, pterygoid. Preorbital width. Post-orbital ,, Orbital Blowhole, width at Zygomatic breadth. Greatest width pmx. Width of braincase across parietals Number of teeth upper R. „ „ „ „ L. Length of tooth row upper R. Hinder end of upper tooth row R. to tip of pmx. L. Number of teeth lower R. Length of lower tooth row R. Hinder end of lower tooth row to tip of mandible Mandible length. Coronoid height. Length of symphysis. Post-temporal length. ,, ,, height. J rostrum length — width at Cranial height. ,, length internal. I 260-6 2 2543 3 246 7 % of parietal width 4567 237-5 253-1 229-3 220-3 «»8 225-5 9 195-7 140-0 139-4 133-3 127-2 137 •(> 121-5 II2-6 116-3 90-8 79 '4 74-3 72 •5 67-4 73 '7 56-9 54-2 74-5 43-3 66 3 61-2 57 •9 58-7 67 'O 48-1 43-4 55-i 30-5 53 •i 52-6 46 '2 47-8 52 5 40-9 41-1 43-9 29-1 34 •8 30-9 3i •6 29-7 36 'X 25-4 23-2 27-5 14-9 181 •8 177-2 170 •8 166-3 177 •8 155-2 141-0 — 116-3 169-1 1 60 •8 157-0 162 '4 147-0 139-9 — 114-9 141 -8 134-9 129 •8 128-2 129 •9 114-9 109-5 120-4 85-1 148 •o 142-3 140 •4 138-0 140 •2 I2I-O 119-1 127-6 95-o 142 •3 135-4 128 •6 135-8 129-9 II3-6 106-5 120-4 87-9 35 •4 32-0 36 •2 33-8 32 9 28-2 28-6 — 29-1 M5 7 140-1 136-9 136-8 140 •2 I2I-0 117-9 — 91-5 54 •9 51-4 53 •2 50-5 52 •I 44-7 48-8 — 41-1 104-0 104-0 107-4 106-9 92-0 94-3 94-9 94-9 87-7 89-5 91-8 89-5 R. L. 95-4 95-4 213-1 48-6 2I-I 47'4 32-0 4O-O 74-3 84-6 206-8 48-6 2O-6 49-1 35-4 42-9 74-9 76-6 94-1 201-8 47-4 22-8 46-2 33'3 33-9 80-4 — 97-9 — — 97-4 — — 104-1 90-6 ~" 103-1 91-7 82-1 85-6 87-6 85-1 85-6 86-4 87-1 87-3 84-8 189-7 90-2 201-0 88-4 180-0 2O-I 44-8 19-6 20-4 50-5 47-9 45-8 35-9 38-0 69-6 27-8 29-8 29-0 69-6 74-7 75-1 — — 59-6 — — 60.3 64-5 82-7 82-1 65-2 — — 66-7 — — 66.7 — — 68-8 42-3 27-4 70-2 75-o 39-8 33-2 68-4 68-9 146-9 34-7 14-9 38-3 22-7 21-3 64-4 80-8 Skull dimensions of P. electro, and their proportions as percentages of parietal width. The numbers at the head of the column refer to the list of specimens on page 177. xl Percentages are to estimated measurement of width of brain case. The caudal aspect of the skull (fig. 5) is dominated by the great extent of the fused occipital bones. The general contour is interrupted by the occipital condyles with the oval foramen magnum between. The general outline is circular, but is inter- rupted by the basioccipital crests, paroccipital crests and notches, nasal boss, zygomatic processes and the posterior margins of the post temporal fossae. Lower ] aw. The lower jaw (fig. 6) is robust, the long axes of the rami making an acute angle with each other where they join at the symphysis, which is short. (The two rami are fused together in this specimen). ELECTRA DOLPHIN 185 The upper and lower margins of the rami are almost parallel up to the hinder end of the tooth row, where they diverge, the upper margin being produced into a low crest between the hindmost tooth and coronoid process. From the coronoid process the margin extends obliquely and rather irregularly to the condyle. Slightly anterior to the level of the mandibular foramina, the profile of the jaw is produced into a keel, which is continued distally into the anterior profile of the jaw. The symphysis is short and in ventral view the prow of the jaw is evenly rounded to each ramus anterior to the keels. FIG. 6. P. electra 1844.10.5.3. type. Lateral view right mandible. Orthographic projection. The tooth rows are again short compared with the length of the jaw, the teeth sitting in discrete alveoli. The proximal teeth are upright in attitude, but pro- ceeding distally they assume an increasingly lateral splay. The crowns of the teeth are separated by a distance of approximately the basal diameter of the teeth. The dimensions of the type skull and of other available specimens are given in Tables i & 2. GROWTH OF THE SKULL (a) Quantitative Discussion. The changes observed with growth are demonstrated quantitatively. The conventional use of condylobasal length as a base for the com- parison of other measurements was discarded in favour of parietal width, following an earlier paper by Fraser and Noble (1968). It is generally accepted that increase in the size of the brain levels off at a com- paratively early stage in the animal's life. The similarity of cranial size in the juvenile and adult indicates conformity to this growth pattern. To determine the rates of growth of the various parts of the skull, values of the various measurements were plotted against their respective parietal widths on a double logarithmic scale. It may be taken that the rates of growth of the various parts, relative to parietal width, satisfy the equation y = bxk. Where x is the parietal width, y the length of the part, b the fractional coefficient (the value of y when x = i), and k the growth coefficient. Calculated values are shown below (Table 3). Values of k over unity indicate an increasing rate of relative growth and those less than unity the converse ; i.e. positive and negative heterogoniety respectively. The skull of the new-born animal is essentially a brain box with diminutive rostrum, supraorbital and zygomatic processes. The growth of the skull is pre- dominantly of these extensions from the brain box. i86 W. H. DAWBIN, B. A. NOBLE, F. C. FRASER TABLE 3 Peponocephala electva Table of growth coefficients for various parts of the skull Growth Reading No. Measurement coefficient 1 Condylo basal length : 1-85 2 Rostrum : length : 2-35 3 Rostrum : basal width : 2-90 4 Rostrum : width 60 mm. anterior to ant-orbital notches : 3-50 5 Rostrum : width at middle : 3-00 9 Preorbital width : 2-60 10 Postorbital length : 2-40 13 Zygomatic breadth : 2-70 15 Width of braincase across parietals : I 33 Rostrum : width at f length : 3-20 34 Cranial height : 1-45 35 Cranial length internal : 0-725 The increase of the condylobasal length is by the growth of two components, rostral and cranial. It will be seen from Table 3 that the contribution of rostral length (2-35) to total length is much greater than is cranial length (0-725). In the rostrum itself growth can be regarded as taking place two-dimensionally, and the table indicates that length, although prominently positively heterogonic is at a lower rate than its lateral expansion. The lateral expansion is not uniform for the whole rostrum. Reading No. 4 is an expression of the lateral expansion that takes place in the basal portion of the rostrum, which is at a lower rate than at the middle of the beak. Towards the tip of the snout (Reading 33) the coefficient indicates a relatively high rate. The interaction of these growth rates is expressed in the shape of the adult rostrum. In the newborn, the preorbital width is considerably less than the postorbital width (5%), and the projections, which develop laterally, are all of less dimension than the parietal width. The reversal of this condition with age is achieved by the faster growth of these parts compared with the expansion of the brain box. Development of the postorbital and zygomatic processes result in the enlargement of the post-temporal fossa. In the new-born, this aperture, through which the temporal muscle passes, is small and subtriangular ; by the growth of the zygomatic and postorbital processes it becomes rounder and is visible in the caudal aspect of the skull of the adult. The coefficients showing development of the brain box (reading 15, 34, 35) indicate a lower rate than for any of the other dimensions considered. But the coefficients of growth throughout the animal's life do not take into account the established fact that brain growth is, in the main, achieved in youth and because of the limitation in available young specimens it is not possible to present coefficients for this rapid phase in the growth of the brain. (b) Topographical Description. Figures 7-11. To demonstrate the changes in the skull with age, four skulls were selected which gave a range of size and development ELECTRA DOLPHIN which could be accepted as indicative of increasing age. follows : — 187 The skulls chosen were as newborn. Reg. No. 1965.6.2.1. Known to be newborn. juvenile. The more complete New South Wales specimen. Chosen on its size and stage of development on generally accepted criteria, such as sutures and occipital crest development. adolescent. 1959.7.9.2. Chosen on its condylobasal length, small size of ossified scapula and non-fusion of vertebral epiphyses. adult. 1844.10.5.3. The type of L. electra Gray. Chosen on condylobasal length and vertebral and scapular development of a skeleton with skull of comparable size and development (W.A.M. No. 4798). General Cranial Shape Cranial Region Newborn : The essential cranial shape is obvious, being smoothly rounded from No. 9* foramen magnum to nasals. The lateral development of supra- orbital processes, maxillary plates, and zygomatic processes is slight ; and the supraoccipital crest is not developed. The greatest width of the skull is between the parietals. Juvenile : The basic shape of the cranium has already become obscured by the No. 7 extension of the supraorbital and zygomatic processes, incipient post-temporal fossa and occipital crest development. The greatest width is between the zygomatic processes. Adolescent : Approximately as in the adult, but the occipital crests are consider- No. 6 ably less developed. Adult : The primary cranial shape is very obscured by the growth of these No. i various features. Sutures Newborn Juvenile All the visible sutures are unfused. The only sutures that are fused are those between the occipital bones ; there is partial fusion of the parietals with the supraoccipital ; the interparietal is fused with the supraoccipital. Adolescent : There is no significant change from the condition in the juvenile. There is indication however of the beginning of fusion of the nasal portion of premaxillae with maxillae. Adult : Nearly every suture has either fused or is showing indication of fusion. The exceptions are : the squamosals which are very loosely joined to the skull ; f rentals with maxillae ; jugals with frontals and maxillae ; nasals with interparietal ; vomer with basisphenoids and basioccipital. * numbers refer to the list on page 177. i88 W. H. DAWBIN, B. A. NOBLE, F. C. FRASER •! PH 03 !-H bo O " JD o Q ELECTRA DOLPHIN 189 (A) (B) (Q (D) 100 mm FIG. 8. Lateral view of skulls of P. electro, ; (A) Newborn, (B) Juvenile, (C) Adolescent, (D) Adult. Orthographic projections. i go W. H. DAWBIN, B. A. NOBLE, F. C. FRASER Development of the mesethmoid The mesethmoid forms the basal septum and posterior wall of the narial passage. Newborn : The ossified mesethmoid is present only as a bony stump projecting posterodorsally into a hiatus between the f rentals. The nasal septum is composed of soft tissue. Juvenile : The mesethmoid is considerably more ossified, covering over the hiatus in the cranium, and it has extended appreciably over the f rentals. The nasal septum is represented by a bony ridge which has not developed to the level of the dorsal surface of the pre- maxillae. Adolescent : The mesethmoid now extends up to make contact with the nasals. The ossified nasal septum is above the level of the lateral borders of the nares. This forward extension however is still restricted and has not yet reached the rostral portion of the skull. Adult : The ossified mesethmoid has extended into the rostrum and is seen as a flattened plate between the premaxillae. Slightly anterior to the level of the premaxillary foramina the septum becomes cartilaginous and continues so to the rostral tip. Maxillary Plate Newborn : The maxillary plates are of reduced extent, their posterior margins being remote from the dorsal margin of the supraoccipital. Their contour anteriorly is much as in the adult but their elevated portions have the contour of the cranium. Juvenile : The maxillae have grown further back towards the occipital crest. Adolescent : As in the adult ; but the maxillae have not extended as near to the supraoccipital crest. Adult : The maxillary plates are spread widely over the f rentals. Rostrally they are roughly horizontal, but curve upwards posteriorly toward the vertex of the skull and supraoccipital crest. Frontal is ex- posed medially between the maxillae and the crests. Occipital Region Newborn : The occipital elements are unfused and two large, posterolateral fontanelles are present between the supraoccipitals and exoccipitals. Each fontanelle, in the prepared specimen, communicates with the foramen magnum by a fissure. On the left the fontanelle is partly occluded by a bony centre ; on the right, this is suggested by the sutural margin. The general outline of the occiput is circular, interrupted by the shallow basicranial groove. The foramen magnum is almost piriform in outline ; its continuity is interrupted by the fissures of the fontanelles and the incomplete fusion ventrally of basioccipital, Juvenile : The supra- and exoccipitals are fused to a great extent. There is a bony insert, which is unfused on the left, in a similar position to ELECTRA DOLPHIN igi Q — PQ, 2 g o II CO O 13 c -!-> a 8 W. H. DAWBIN, B. A. NOBLE, F. C. FRASER centrum. Incipient transverse processes stem from the neural arches of the first 17 vertebrae, from which point caudally they can be distinguished on the centra, diminishing to disappearance in the posterior caudal region. Rudimentary superior transverse pro- cesses are present on the vertebral arches of the 4th — 7th cervical vertebrae, but there are no indications of inferior transverse processes. Adolescent : The atlas, axis and third cervical vertebra of the adolescent are fused ; the atlas and axis being joined at the neural spine and base of the centrum. There is still separation between the neural arches of the atlas and axis dorsolaterally, affording a passage for the second cervical spinal nerve. The third cervical vertebra is fused to the posterior face of the axis, though the limits of the two centra can be discerned. The neural spines of all three vertebrae are fused with each other. The third cervical has an obvious upper transverse process and rudimentary lower transverse process. The vertebrae are at an advanced state of development ; the verte- bral arches are fused to their centra, the transverse processes are well developed. None of the epiphyses of the centra is fused to its centrum. The last 10 vertebrae are missing (from a total of 82 indicated by Nakajima & Nishiwaki, 1965). Neural arches extend tailwards to the 59th vertebra. Adult : Ventrally, ankylosis of the atlas, axis and third vertebra is more extensive than in the adolescent, but dorsolaterally the limits of the centra of the third cervical vertebra and the axis are still traceable. The neural spine of the axis, which in the adolescent is bluntly pointed, has developed in the adult into a robust spine, terminally bifurcate. The arch of the vertebra is not fused to that of the axis and is incomplete apically. The transverse processes of the axis are rounder, more pointed and extending further laterally than in the adolescent. The series of vertebrae is not complete, C6 is missing and there are gaps in lumbar and caudal successions, the terminal caudals are all absent. Nevertheless the vertebrae remaining indicate by the absence of unfused epiphyses that the animal was physically mature. The apex of the anticline of the neural spines is at about the 30th vertebra in both adolescent and adult . Post-zygapophyses are present to about the I5th vertebra. The pre-zygapophyses are present on the cervical and anterior thoracic vertebrae, but they are gradually replaced by the increasing prominence of the meta- pophyses in the anterior thoracic region. The metapophyses in their turn gradually decrease in prominence in the anterior lumbar region, but behind the anticlinal apex they are strongly developed up to the point where the vertebral arches begin to disappear. ELECTRA DOLPHIN 199 Rib Neck Vestiges The existence of rib neck vestiges on transverse processes has been noted by Flower (1872), Slijper (1936) and Fraser (1940). Slijper notes the condition in L. albirostris so far as the genus Lagenorhynchus is concerned. The presence of these rib neck vestiges is associated with a change in the attachment of the ribs to the vertebral column. In the adult and adolescent electra there are five and six ribs respectively which possess both capitulum and tubercle. The remaining ribs have neither neck nor capitulum. This change from complete to ' tubercular ' ribs is clearly defined on the ribs themselves, but on the associated vertebrae it is expressed as a transitional series, so that if the number of complete ribs is five (as in the adult), the vertebra associated with the sixth rib has attached to it a capitular portion differing very little from that of the preceding rib, except that it is fused to the transverse process and separate from the rib (fig. 13). The heads of the vestiges are in very close proximity to the articular facets on the fifth vertebra, and themselves bear articular facets. The vestige is missing on this vertebra in the adolescent, which has six complete ribs. FIG. 13. Anterolateral view of thoracic vertebrae 5-8 of adult, to show the rib neck vestiges. On the transverse processes of the seventh thoracic vertebra of both the adult and adolescent, there are bilateral, short, hook-like processes whose apices point in the direction the necks of the ribs would follow if present, toward vestigial catapophyses on the vertebra in front. The vestiges on the eighth vertebra of both the adult and adolescent are low, barely perceptible eminences on the ventral surfaces of the transverse processes. There are very reduced catapophyses on the vertebra in front. The articular facets for the tubercles of the ribs on the sixth and eighth, and fifth and eighth thoracic vertebrae in the adolescent and adult respectively, are oval, while those of the intervening vertebrae are sub-triangular, implying the incorporation of additional bony elements. In terrestrial mammals, the majority of the ribs have distinct tubercle and 200 W. H. DAWBIN, B. A. NOBLE, F. C. FRASER capitulum. The cetaceans are distinguished by the increased number of ribs having single articulation with the corresponding vertebra. The odontocetes have three patterns of vertebra-rib connection ; anteriorly, a double connection at the head and tubercle with the centrum and transverse process respectively ; further back, a tuberculo-transverse process articulation only ; caudally, there are rib elements which do not have an obvious articulation with the vertebral column. It may be noted further that in odontocetes, the sternal ribs are completely ossified. The form of double articulation of the ribs with the vertebral column, and possess- ion of ossified sternal ribs anteriorly, indicate the rigidity of the fore part of the rib basket. Posteriorly, the thoracic cage, being without capitular-centrum connections of the ribs and having a much looser association with the sternum, would allow greater flexibility of the body, as pointed out by Slijper. The connection of the head to trunk demands a strong framework for the relevant musculature, part arising from the vertebral column, part from the ribs, and part from both. Further, the attachment of the pectoral limbs is dependent on muscles arising from anterior vertebrae and ribs, again demanding stability of the rib cage for their proper functioning. The locomotion of these animals requires a flexibility of the vertebral column and development of axial musculature. This flexibility is initiated in the posterior thoracic region and is achieved by reduction of the vertebral apophyses and develop- ment of the inter-vertebral discs ; it seems likely that the removal of the capitular articulation in this region in the vertebral column contributes to this flexibility. The enlargement of the axial musculature required in connection with cetacean locomotion necessitates a complementary extension of neural spines and transverse processes. In the individual dolphin the progressive lateral extension of the trans- verse processes of the thoracic vertebrae is unavoidably associated with increasing attenuation of the rib neck. It seems reasonable that a point is reached where the mechanical stability of the neck is lost, resulting in ribs possessing tubercles only. Summary Two specimens of dolphin stranded on the coast of New South Wales are identified as belonging to the species Peponocephala electra. As Gray's description of the holotype of Lagenorhynchus electra is brief, a redescription is given, together with orthographic projections of various aspects of the skull. Ten skulls available for comparison have been measured and the data applied to quantitative assessment of the skull. A topographical survey was made of four skulls of increasing age to show the changes in skull form from birth to physical maturity. Comparative notes have been made of some of the alterations in the axial skeleton with age. ACKNOWLEDGMENTS The authors are indebted to Professor Kenneth Norris, University of California, for making available the skeleton of the newborn specimen, Dr. D. L. Ride, Director of the Western Australian Museum for lending the skeleton W.A.M. 4798, and to Dr. R. G. Van Gelder, American Museum of Natural History, New York, for the loan of the specimen A.M.N.H. 4300. ELECTRA DOLPHIN 201 Thanks are due to the many people who wrote or sent photographs relating to the Crowdy Heads school, especially Mrs. W. J. Ward, Mr. H. Anderson and Mr. L. Elford. REFERENCES DAWBIN, W. H. 1963. Mass stranding of dolphins Lagenorhynchus sp. Bull. Aust. mamm. Soc. 6 : 14. FLOWER, W. H. 1876. An introduction to the Osteology of the Mammalia, and ed. Revised. 1-344. London (Macmillan & Co.) FRASER, F. C. 1940. Three anomalous dolphins from Blacksod Bay, Ireland. Proc. R.Ir.A. 45 : B, 17 : 413-455. & PURVES, P. E. 1960. Hearing in Cetaceans — II Evolution of the accessory air sacs and the structure and function of the outer and middle ear in Recent Cetaceans. Bull. Brit. Mus. (N.H.), 7 : No. i, 1-140. & NOBLE, B. A. 1968. Skull of Lagenorhynchus cruciger from Livingston Island, South Shetland Islands. Bull. Br. Antart. Surv. 15 : 29-38. GOODWIN, G. G. 1945. Record of a porpoise new to the Atlantic. /. Mammal. 26 : 195. GRAY, J. E. 1846. Zoology of the Voyage of H.M.S. Erebus and Terror . . . 1839-43. 1 (Mammalia) : 13-53. HOWELL, A. BRAZIER 1930. Aquatic Mammals. 1-338. Baltimore (Charles C. Thomas). NAKAJIMA, M. & NISHIWAKI, M. 1965. The first occurrence of a porpoise (Electra electra) in Japan. Sci. Rep. Whales Res. Inst. 19 : 91-104. NISHIWAKI, M. & NORRIS, K. 1965. A new genus, Peponocephala, for the odontocete cetacean species Electra electra. Sci. Rep. Whales Res. Inst. Tokyo. 20 : 95-99. OWEN, R. 1866. On some Indian cetacea collected by Walter Elliott, Esq. Trans, zool. Soc. Lond. 6 : 17-47. PEALE, T. R. 1848. United States Exploring Expedition during the years 1838-1842, under the command of Charles Wilkes U.S.N. 8 (Mammalogy & Ornithology) : xxv+ 17-388. Philadelphia. SLIJPER, E. J. 1962. Whales. 1-475. London (Hutchinson) . TRUE, F. W. 1889. Contributions to the natural history of the cetaceans. A review of the family Delphinidae. Bull. U.S. natn. Mus. 36 : 1-192. WILLIAM H. DAWBIN, D.Sc. SCHOOL OF BIOLOGICAL SCIENCES UNIVERSITY OF SYDNEY N.S.W., AUSTRALIA BRUCE A. NOBLE, B.Sc. c/o BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON, S.W.7 FRANCIS C. FRASER, C.B.E., D.Sc., F.R.S. c/o BRITISH MUSEUM (NATURAL HISTORY) CROMWELL ROAD LONDON, S.W.7 Printed in Great Britain by Alden & Mowbray Ltd at the Alden Press, Oxford THE SPECIES OF MACROPHTHALMUS (CRUSTACEA: BRACHYURA) IN THE COLLECTIONS OF THE BRITISH MUSEUM (NATURAL HISTORY) r \ 1-5 JAN 1971 I V5> R. S. K. BARNES BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 20 No. 7 LONDON : 1970 THE SPECIES OF MACROPHTHALMUS (CRUSTACEA: BRACHYURA) IN THE COLLECTIONS OF THE BRITISH MUSEUM (NATURAL HISTORY) BY RICHARD STEPHEN KENT BARNES Department of Zoology, University of Bristol Pp. 203-251 ; 10 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 20 No. 7 LONDON : 1970 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, is issued in Jive series corresponding to the Departments of the Museum, and an Historical series. Parts will appear at irregular intervals as they become ready. Volumes will contain about three to four hundred pages, and will not necessarily be completed within one calendar year. In 1965 a separate supplementary series of longer papers was instituted, numbered serially for each Department. This paper is Vol. 20, No. 7 of the Zoological series. The abbreviated titles of periodicals cited follow those of the World List of Scientific Periodicals. World List abbreviation Bull. Br. Mus. nat. Hist. (Zool.). Trustees of the British Museum (Natural History), 1970 TRUSTEES OF THE BRITISH MUSEUM (NATURAL HISTORY) Issued 28 December, 1970 Price £1.40 THE SPECIES OF MACROPHTHALMUS (CRUSTACEA: BRACHYURA) IN THE COLLECTIONS OF THE BRITISH MUSEUM (NATURAL HISTORY) By R. S. K. BARNES CONTENTS Page 206 207 207 208 211 2I4 2I4 216 219 219 220 222 223 . . . . . 225 226 226 228 229 232 232 232 236 C. SUBGENUS Venitus ......... 236 1. M. latreillei 236 2. M. pectinipes ......... 237 D. SUBGENUS Mopsocarcinus . . . . . . . .241 1. M. bosci .......... 241 2. M. punctulatus ......... 242 E. SUBGENUS Hemiplax ......... 242 i. M. hirtipes .......... 242 F. SUBGENUS Tasmanoplax ........ 243 i. M. latifrons ......... 243 DISCUSSION ........... 243 CONCLUSIONS ........... 246 ACKNOWLEDGEMENTS ......... 247 REFERENCES ....... 247 INTRODUCTION A. SUBGENUS Macrophthalmus i. M. brevis 2. M. transversus 3- M. parvimanus 4- M. dilatatus . a. M. d. dilatatus b. M. d. sulcatus 5- M. telescopicus 6. M. crassipes 7- M. laevimanus 8. M. convexus , 9- M. grandidieri . 10. M. graeffei B. SUBGENUS Mareotis . i. M. depressus 2. M. japonicus . 3- M. tomentosus 4- Af. definitus 5- M. pacificus . - 6. M. erato . 7- M. crinitus . 206 R. S. K. BARNES SYNOPSIS The British Museum holds twenty three species of the genus Macrophthalmus (Crustacea : Brachyura). Nine of these, M. depressus, M. dilatatus, M. erato, M. grandidieri, M. laevimanus, M. parvimanus, M. pectinipes, M. tomentosus & M. transversus, are redescribed and figured. Material from the Royal Society's 1967-1968 Expedition to Aldabra is also included. The convergence with certain sesarmine grapsids displayed by M. hirtipes, clinal changes in mor- phology in M . dilatatus, and the systematic value of a number of features of the genus are described and discussed, in addition to a review of the B.M. collection. INTRODUCTION THE following is a review of the British Museum collection of crabs of the genus Macrophthalmus Latreille (Ocypodidae : Macrophthalminae), it being the fifth con- tribution towards an eventual revision of this genus (see Barnes, ig66a. ; ig66b ; 1967 ; ig68a). The review follows the same basic pattern as that of an earlier paper dealing with the species of this genus from Australia and adjacent regions (Barnes, 1967), and here only those species not covered by that publication will be fully described and figured, i.e. those marked by an asterisk in the list given below. Wherever possible, regression equations have been calculated to show the changes in the carapace length/breadth ratio and that of breadth of front/carapace breadth with changes in size (see Barnes, igGSb). The collection contains material previously described and discussed by de Haan (1835), Adams & White (1848), Miers (1884 ; 1886), Lanchester (igooa ; igoob), Laurie (1906 ; 1915), Caiman (1927), Gordon (1931), Tweedie (1937) and McNeill (1968). Here, the two hundred and forty four specimens comprising the collected material are assigned to the following twenty three species : M. bosci Audouin ..... 52 specimens ? M. brevis (Herbst) . . . . . 16 ,, M. convexus Stimpson . . . . n ,, M. crassipes H. M. Edwards . . . 8 ,, M. crinitus Rathbun . . . . . 8 ,, M. definitus Adams & White . . . 6 ,, *M. depressus Riippell . . . . . 27 ,, *M. dilatatus (de Haan) . . . . 8 „ *M. erato de Man ..... 9 ,, M. graeffei A. M. Edwards . . . . 3 *M. grandidieri A. M. Edwards 10 ,, M. hirtipes (Jacquinot) . . . . 8 M. japonicus (de Haan) . . . . 10 ,, *M. laevimanus H. M. Edwards . . . i ,, M. latifrons Haswell . . . . . 2 M. latreillei (Desmarest) .... 22 „ M. pacificus Dana . . . . . 9 ,, *M. parvimanus Guerin . . . . I1 ,, *M. pectinipes Guerin . . . . . 10 ,, 1 Also included in this report are 13 specimens of M. parvimanus collected by the Royal Society's 1967-1968 Expedition to Aldabra. SPECIES OF MACROPHTHALMUS 207 M. punctulatus Miers i specimens M. telescopicus (Owen) . . . . 13 ,, *M. tomentosus Souleyet . . . . 4 ,, *M. transversus (Latreille) . . . . 5 ,, The collection therefore contains approximately two thirds of the probably valid species included in this genus. With respect to the validity, or otherwise, of a number of published species, the author has attempted to keep the majority of such discussion for a proposed full revision of Macrophthalmus. However, some com- ments have been made on a number of new synonymies which have become apparent during this study. One particularly difficult case to unravel at this juncture is centred around M. brevis (= M. carinimanus H. M. Edwards). A considerable number of minor variations and different geographical populations of this species have almost certainly been made the basis for a plethora of specific names, but an adequate treatment of this species must await the examination of more material from more localities. That part of the problem which impinges on M. dilatatus has, however, been dealt with in some detail. Other difficult cases are centred around M. telescopicus and M. latreillei. In addition, the status of the following species has been deliberately excluded from discussion in this report — M. consobrinus, M. graeffei, M. latipes, M. milloti and M. teschi. Any modern worker studying Macrophthalmus species from a large geographical area must very quickly become aware that the characters displayed by these organ- isms do not remain constant, but vary (a) with size of the animal, (b) amongst material from a single locality, and (c) from locality to locality. But as a result of the patchiness of much of the collecting which has produced the specimens examined, our knowledge of many species over much of their range is negligible. In a few cases clinal changes in morphology can be discerned (see M. dilatatus) and in more cases intermediate forms between two distinct structural types can be found. But often samples are inadequate for more than tentative judgements. Complete synonymies are given only for those species not covered by Barnes (1967). The dimensions given under " Material examined " headings below are of greatest carapace breadth. A. Subgenus MACROPHTHALMUS Sensu stricto i. Macrophthalmus (Macrophthalmus) brevis (Herbst, 1804) Foreword This is an extremely problematical species, with an intensely confused synonymy, but since very few specimens have been seen by the author, it would be premature to do more here than indicate the nature and magnitude of the problem. Six species, M. laevimanus H. M. Edwards, M. sandakani Rathbun, M. simdentatus Shen, M . malaccensis Tweedie, M. malayensis Tweedie and M. travancorensis Pillai, have been described from the area between the Gulf of Manaar and Hainan Island. 208 R. S. K. BARNES Each is known only from one or two localities and between them they total only about twenty specimens. Further, none of these authors refers to any of the other authors' species. Secondly, there are two very poorly known species, M. sulcatus H. M. Edwards and M. brevis (Herbst), known from approximately the same area. These eight species, judging from the published descriptions, all appear to grade into one another in a non-linear manner, and there are additional links to a ninth species, M. dilatatus (de Haan). Lastly, Lanchester's (igooa) record of M. crassipes H. M. Edwards and Tweedie's (1937) record of M . c.f. crassipes are not of that species, but together with Lanchester's M . dilatatus carens form a tenth unit related to the previous nine. All the material of ' M. brevis ' in the B.M. collection falls into this latter, tenth unit. The status of M. sulcatus, M. sandakani and M. malaccensis will be discussed under the section on M. dilatatus, and M. laevimanus and M. malayensis are considered in a separate section below. MATERIAL EXAMINED. 8 $$ (12-0-17-0 mm), 8 $ $ (11-8-19-7 mm). B.M. Reg. Nos-82.24, 1900.10.22.277-284 (Syntypes of M. dilatatus carens}, 1900.10.22.285-292. LOCATIONS. Singapore, Malacca. COMMENTS. The material with the registration number of 82.24 is that collected by the " Alert " and identified by Miers as M. dilatatus (it is not recorded in the results of the " Alert " voyage — Miers, 1884 — presumably as a result of an oversight) ; Lanchester (igooa) regarded it as being identical with his M. dilatatus carens. 1900.10.22.277-284 is the type material of the latter subspecies (Lanchester, igooa), which was considered by Tweedie (1937) to be a series of juvenile M. brevis. That with the number of 1900.10.22,285-292 was recorded by Lanchester (igooa) as M. crassipes, and Tweedie was of the opinion that, together with his own M. c.f. crassipes material, it might form a new subspecies of M. crassipes (Tweedie, 1937). All these specimens are juveniles, which increases the uncertainty of identification. 82.24 and 1900.10.22.277-284 are almost certainly of the same species, to which 1900.10.22.285-292 may belong, but there are a number of noticeable differences between the latter and the two former. What perhaps contributes most to their incertae sedis position is that, although juvenile, they are large and possess carapaces relatively longer than would be expected for their size. For example, the largest specimen (19-7 mm) has a carapace length one millimetre larger than that of a " M. malaccensis " of carapace breadth 21-8 mm. 2. Macrophthalmus (Macrophthalmus) transversus (Latreille, 1817) Gonoplax transversus Latreille, 1817 Macrophthalmus transversus : Latreille, 1829 ; H. M. Edwards, 1837 '• H. M. Edwards, 1852 ; Cano, 1889 ; deMan, 1892 ; Tesch, 1915 ; Kemp, 1919 MATERIAL EXAMINED. 4 ^ (17-0-23-0 mm), i $ (18-7 mm). B.M. Reg. Nos— 1919.11.91-94, 1951.4.19.2. LOCATIONS. Chandpur, Balasore (India). DESCRIPTION. Front deflexed ; markedly constricted between bases of ocular SPECIES OF MACROPHTHALMUS 209 peduncles ; smooth margined ; with almost straight anterior margin and faint median furrow. Upper orbital border strongly curved, transversely directed ; margin studded with rounded granules increasing in size towards external orbital angle, granules nearest to that angle large and tubercular. Lower orbital border serrated by large, curved, pointed tubercles ; with from three to six very large, more or less flattened spines in centre (each approx. three times longer than normal tubercle and nine times the surface area). FIG. i. M. transversus — a, anterolateral carapace teeth and ocular peduncle, b, left male chela (outer surface), c, external margin of ischium of external maxilliped. Scale lines a & c i mm., b. i cm. Three well defined anterolateral teeth (see Fig. la). External orbital angle large, elongate, strongly pointed, directed straight outwards ; anterior margin with con- tinuation of granulation of upper orbital border ; lower margin with rounded granules ; tip often formed by elongate tubercle ; separated from second lateral tooth by deep, narrow incision, wider and more U-shaped in juveniles. Second lateral tooth large, of the shape shown by Fig. la, directed outwards and forwards ; anterior margin with rounded granules ; convex or straight outer margin with three large tubercles (in adults), the largest anteriorly forming point of tooth, second in centre of margin, third at posterior angle directed posteriorly ; separated from third lateral tooth by deep, narrow incision. Third lateral tooth moderately large, triangular, pointed, 210 R. S. K. BARNES directed outwards and slightly forwards, with large tubercle at tip ; margins straight, outer margin smoothly continuous with lateral carapace margin. Carapace completely covered with small, rounded granules ; with well denned, deep furrows ; with well defined, raised clumps of granules on branchial regions, each bearing one very large tubercle in its centre, accessory granules of ten lacking, " clump" then represented only by tubercle, additional similar tubercle often present between positions of first and second clumps (tubercle formed from second clump) ; with row of four to six smaller, pointed tubercles between branchial clumps or tubercles and lateral carapace margins ; with abruptly sloping sides ; without surface hair. Lateral margins posteriorly convergent, with row of about eight large tubercles on margin, evenly spaced from third lateral tooth to posterior angle of carapace, tubercle at that angle particularly large. With row of granules near to and subparallel with posterior margin. Greatest carapace breadth across external orbital angles. Female carapace without granules excepting on branchial and lateral regions. Ocular peduncles very long and narrow, projecting beyond external orbital angle for between one tenth and one fifth of their length. Male cheliped. (a) Merus. Upper margin with scattered granules and from one to three large spines centrally; inner margin with row of tubercles along length, of which any number from two to six may be converted into very long spines (often differentially on the two meri, e.g. one with six, the other three) ; outer margin with scattered granules and one or two large spines distally. Inner surface without granules, with patch of hair centrally and proximally near inner margin ; outer surface with scattered granules near upper and outer margins ; lower surface with scattered granules near outer margin and thickish hair near inner margin. (b) Carpus. Almost hairless. Outer surface with scattered granules, those near lower margin large and tuberculiform ; inner surface more or less smooth, except for a large spine near upper margin and similar spine in central region directed towards palm. Upper margin with scattered small granules ; lower margin with irregular tubercular granules. (c) Palm. Very elongate. Outer surface closely covered with small, pointed granules ; with prominent longitudinal ridge bearing a row of small, pointed tubercles along crest ; anterior margin without notch. Inner surface closely covered with granules, with thick hair over whole surface, with exception of extreme proximal region and area near lower margin, with large spine, and accessory tubercles in large specimens, near to and directed towards carpus. Upper margin with row of from four to eight large spines, increasing in size towards carpus, with granules between spines ; lower margin with pointed granules as on outer surface (see Fig. ib). (d) Index. Markedly deflexed, elongate, very thin. Outer surface with small, sparse granules, with faint, feebly granular continuation of longitudinal ridge of palm, granules decreasing in size distally ; inner surface sparsely granular, with dense hair near cutting margin. Cutting margin with very large, tall, laterally flattened, pointed tooth in centre, tooth either spiniform with straight entire margins, or with a crenulated posterior margin (in some large specimens, the tooth may take the form of two divergent spines joined together at their base), a second smaller spiniform tooth, with entire margins, situated half way between larger tooth and tip of index, re- SPECIES OF MACROPHTHALMUS 211 mainder of margin usually without further conspicuous granules ; lower margin with pointed granules and few tubercles. (e) Dactylus. Markedly deflexed, elongate, very thin. Outer surface closely covered with small granules ; inner surface completely covered by mat of thick hair. Upper margin with small, pointed granules ; cutting margin with large, triangular, spiniform or crenulated tooth near base, with irregular number of small, cylindrical spines in distal half, with small, irregular granules and tubercles over remainder. Extreme tip of dactylus hooked. Pereiopod meri with from one to six large spines on the distal part of posterior lower margin in large specimens. Male abdomen with bulge in morphologically anterior half of lateral margins of sixth segment. External maxilliped. Internal margin of ischium almost straight or slightly convex ; external margin with a marked and precise concavity proximally (see Fig. ic), distal portion of margin smoothly but slightly convex. Internal margin of merus convex ; external margin smoothly convex, without differentiated con- vexities ; anterior margin shallowly concave. First male pleopod almost straight, without well developed terminal process, without hair on internal margin except at tip. Central convexity of epistome small and pointed. DIMENSIONS. Over the size range examined, the carapace length/breadth ratio varies from 0-40 : i to 0-44 : i, and the breadth of front /carapace breadth ratio from 0-12 : i to 0-13 : i. COMMENTS. M. transversus, the type species of the genus, is remarkable for the extremely thin chelae of both sexes (the female chela is as thin as that of the males and possesses similar spiniform teeth, but is otherwise not spiniferous) and for the extremely tubercular and spiniferous carapace, chelipeds and pereiopods, it being rivalled in the latter only by M. pectinipes. As pointed out by Kemp (1919), the additional joint on the dactylus of the male cheliped, as described and figured by Tesch (1915), is the result of a partial fracture of the base of that finger in his specimen. 3. Macrophthalmus (Macrophthalmus) parvimanus Guerin, 1834 Macrophthalmus parvimanus Guerin, 1834 : H- M- Edwards, 1837 ; H. M. Edwards, 1852 ; Richters, 1880 ; Miers, 1884 ; Balss, 1934 '< Taylor, 1968 Macrophthalmus convexus kempi Gravely, 1927 Macrophthalmus convexus : Kemp, 1919 (part) Macrophthalmus consobrinus : Crosnier, 1965 MATERIAL EXAMINED. 7 ^ (8-1-24-9 mm), 7 ?? (10-8-23-3 ™m)- B-M- Reg. No. 82.24, and Royal Society Expedition, Aldabra 1967-1968 (" Lagoon mud, Dune Jean Louis Creek, Aldabra, i8/xi/i967 "). LOCATIONS. Mahe (Kerala), Aldabra. DESCRIPTION. Front deflexed ; markedly constricted between bases of ocular 212 R. S. K. BARNES peduncles ; smooth margined ; with straight or slightly bilobed anterior margin ; with shallow median furrow. Upper orbital border curved, markedly backwardly sloping ; margin beaded by small, rounded granules. Lower orbital border serrated by large, evenly spaced, tubercular granules along entire length. Two well defined and one very poorly defined anterolateral teeth (see Fig. 2a), beaded by small granules along all margins. External orbital angle large, pointed, directed outwards and forwards at its tip ; separated from much smaller second lateral tooth by wide, V-shaped or very narrow incision. Second lateral tooth small, pointed, triangular, directed straight outwards ; separated from third lateral tooth by very small incision. Third lateral tooth very small or absent. FIG. 2. M . parvimanus — a, anterolateral carapace teeth, b, left male chela (outer surface). Scale lines — 5 mm. Carapace smooth and shiny to naked eye (except for granular clumps on branchial regions), lateral areas microscopically granular ; with faint, shallow furrows, except- ing well developed circumgastric ; with well developed granular clumps on branchial regions ; lateral borders with mat of short, fine hair. Greatest carapace breadth across external orbital angles. Lateral margins markedly convergent posteriorly, with rows of long, silky hairs along their length. Ocular peduncles long and narrow, cornea extending as far as, or slightly beyond, tip of external orbital angle. Male cheliped. Unique in that it is not sexually dimorphic in this species ; males with small, slender, weak chelae of the same pattern as found in the females of other Macrophthalmus species. (a) Merus. Upper and inner margins with long, fine hair ; outer margin with row of very small granules. All surfaces without granules ; scattered hairs only on inner surface. SPECIES OF MACROPHTHALMUS 213 (b) Carpus. Without granules or tubercles. Upper margin with fringe of long hairs ; lower margin with few scattered hairs ; outer anterior margin with long hair mainly in lower portion. Both surfaces smooth. (c) Palm. Outer surface finely granular, with longitudinal ridge very close to lower margin ; inner surface without granules, with square or rectangular patch of short thick hair centrally, with row of long, fine hairs near to and subparallel with upper margin. Upper margin with row of small granules and row of long, fine hairs ; lower margin with small granules on the longitudinal ridge (see Fig. 2b) . (d) Index. Undeflexed. Outer surface smooth, except for marked, agranular continuation of longitudinal ridge near lower margin, and row of long hairs near distal cutting margin ; inner surface smooth, except for row of long hairs near distal cutting margin. Cutting margin without differentiated tooth except in large specimens, in which from eight to ten granules are associated to form a long, low tooth, from one to two fifths of the length of the margin away from its base, with row of rounded granules proximally ; lower margin smooth. (e) Dactylus. Straight. Outer and inner surfaces smooth, apart from row of hairs near distal cutting margin. Cutting margin without differentiated tooth, except in large specimens in which five or six granules near the base are associated to form a distinct tooth, with few small granules proximally and centrally ; upper margin with fringe of long, fine hair. Pereiopod meri, carpi and propodi with quite heavily granular surfaces and margins, upper margin of merus with row of hairs, and small subterminal spine. Male abdomen. Lateral margins of sixth segment with bulge in morphologically anterior position. Sternal surfaces granular. External maxilliped. Internal and external margins of ischium more or less straight through much of their length, distally convergent. Internal margin of merus convex ; external margin with marked posteroexternal convexity and faint anteroexternal convexity ; anterior margin with shallow concavity. First male pleopod with very well developed terminal process, without hair on internal margin. DIMENSIONS. Carapace length = 0-46 carapace breath + 0-82 (Standard devia- tion 0-21), Breadth of front = 0-088 carapace breadth + 0-59 (Standard deviation 0-05). COMMENTS. This species was shrouded in obscurity until the publication of Balss's (1934) paper, in which he showed that there was indeed a species of Macro- phthalmus with juvenile-like adult male chelae. Previously most authors had accepted Tesch's (1915) contention that M. parvimanus was probably based on a female specimen of Uca. M. parvimanus is, in fact, extremely closely related to the well known M. convexus, the only character separating the two species being the peculiar cheliped of the former species. The large (32-5 mm), abnormal male recorded by Kemp (1919) from the " upper end of the Gulf of Manaar " is clearly a specimen of M. parvimanus. Kemp noted that " The chela differs from that of the female [of M. convexus] in only two points, — in the possession of rudimentary teeth on the fingers and in the hairy 214 R- S. K. BARNES covering of the inner surface " and that " In all other respects the specimen agrees precisely with normal examples of the species [M. convexus] " (Kemp, 1919 : 389 and see his Plate 24, fig. 2). The specimens recorded and described by Gravely (1927) as M. convexus kempi, from the Gulf of Manaar, also belong to this species. Of his specimens, Gravely (1927 : 150) states " As . . . the seven males (two small) in our collection all agree with this abnormal specimen [Kemp's] it is evident that they represent a distinct local race for which I propose the name kempi. " Besides the Gulf of Manaar, however, this species is known from the Seychelles (the type locality) and nearby islands, and from Madagascar, since Crosnier's (1965) record of M. consobrinus was based on material of M. parvimanus. This was ascertained by the examination of part of Crosnier's material in the Museum National d'Histoire Naturelle, Paris, through the courtesy of Dr. D. Guinot. 4. Macrophthalmus (Macrophthalmus) dilatatus (de Haan, 1835) In this paper, M. dilatatus (sensu de Haan and subsequent authors) is regarded as the nominal subspecies of M. dilatatus (sensu novo) and M . sulcatus, M. sandakani and M. malaccensis are regarded as being synonymous and forming a second sub- species M. dilatatus sulcatus H. M. Edwards, 1852 (comb. nov.}. a. M. dilatatus dilatatus (de Haan, 1835) Ocypode dilatata de Haan, 1835 Macrophthalmus dilatatus : H. M. Edwards, 1852 ; de Man, 1890 ; Ortmann, i8Q4a ; Ortmann, 1897 ; ? Doflein, 1904 ; Tesch, 1915 ; Sakai, 1934 > Sakai, 1939 ; Sakai, 1965 nee Macrophthalmus dilatatus car ens Lanchester, igooa MATERIAL EXAMINED. 2 $$ (26-3 & 30-8 mm), i $ (31-5 mm). B.M. Reg. Nos— 1935.3.19.37-38, 1961.6.5.92. LOCATIONS. North China, Tokyo Bay or Sagami Bay (Japan). DESCRIPTION. Front deflexed ; constricted between bases of ocular peduncles ; with smooth margins, slightly bilobed or straight anterior margin, shallow median furrow. Upper orbital border markedly curved and almost transversely directed, with little backwards slope ; margin beaded by small, rounded granules. Lower orbital border serrated by large, widely spaced, tubercular granules, often with smaller granules alternating with the large. Two well defined and one poorly defined anterolateral teeth (see Fig. 3a). Ex- ternal orbital angle narrow, elongate, pointed, directed outwards and forwards to a variable degree (varying from straight outwards to outwards and up to 20° forwards, measured from the transverse carapace axis) ; anterior margin with small granules continuous with those on upper orbital border ; posterior margin smooth ; separated from second lateral tooth by deep, but narrow incision. Second lateral tooth large, wedge-shaped, directed outwards and forwards ; anterior margin smooth or with small granules ; posterior margin straight or convex, with granules ; tip extends as SPECIES OF MACROPHTHALMUS 215 far as, or slightly beyond, that of external orbital angle ; separated from third lateral tooth by narrow incision. Third lateral tooth variable in size, directed outwards and forwards ; with granular margins. Carapace of darkish colour, covered with medium sized granules to a variable extent, with central areas almost devoid of granules in some specimens ; with well denned, deep furrows ; with distinct raised clumps of tubercular granules on branchial regions ; with abruptly sloping sides. Greatest carapace breadth across external orbital angles and second lateral teeth, or across latter alone. Lateral margins posteriorly convergent, with rows of hair along length ; posterior margin granular. Ocular peduncles long and narrow, cornea extending to tip of external orbital angle. FIG. 3. M. dilatatus dilatatus — a, anterolateral carapace teeth, b, right male chela (outer surface). Scale lines — i cm. Male cheliped. (a) Merus. Upper margin with large tubercle centrally and patch of hair proximally ; inner margin densely haired, with one or two very large tubercles distally ; outer margin granular, with one or two large tubercles distally. Lower surface sparsely granular, with dense mat of hair near and continuous with that on inner margin, hair may extend over most of surface ; inner surface more or less smooth, with hair near inner margin ; outer surface with sparse hair near upper margin and sparse granules near outer margin. (b) Carpus. Almost hairless. Outer surface smooth centrally, with scattered granules near upper and lower margins ; inner surface smooth, except for large tubercle dorsally and similar tubercle centrally near joint with palm. Lower margin with few, large granules distally ; upper margin with sparse row of granules distally. (c) Palm. Elongate. Outer surface with very large, evenly spaced, rounded 216 R. S. K. BARNES tubercles in upper half, the lowest tubercles largest and forming a row (see Fig. 3b), area between row of tubercles and longitudinal ridge smooth, longitudinal ridge prominent and with large granules on crest decreasing in size towards index, area below ridge heavily granular, anterior margin with deep notch ; inner surface heavily haired over all but lower and proximal region, boundary to haired portion marked by row of granules and large spine near to and directed towards carpus, lower proximal region heavily granular, especially near lower margin. Upper margin with row of large tubercles, largest centrally ; lower margin granular. (d) Index. Markedly deflexed in adults. Outer surface more or less smooth, except for low, agranular continuation of longitudinal ridge ; inner surface heavily haired near cutting margin, with small granules near lower margin. Cutting margin with long, low, crenulated tooth occupying proximal half to three fifths, with a few large granules distally in adults ; lower margin granular. In adults, index strongly curved near base, correlated with interdigital notch. (e) Dactylus. Slightly curved, oriented almost vertically in adults. Outer surface smooth apart from row of granules near upper margin ; inner surface heavily haired (hair on inner surfaces of index and dactylus continuous with that on palm) . Upper margin granular ; cutting margin with low tooth formed from four or five contiguous granules in a line at base, with rounded granules and dense hair along remainder. Base of cutting margins of index and dactylus widely separate. Pereiopod men with hair and small subterminal spine on upper margin, without conspicuous granules. Male abdomen. Lateral margins of sixth segment with bulge in morphologically anterior half. External maxilliped. Internal margin of ischium convex ; external margin more or less straight. Internal margin of merus convex ; external margin curving smoothly into anterior margin, without distinct posteroexternal convexity ; anterior margin smoothly concave. First male pleopod curved, with well developed terminal process, without hair on internal margin except at tip. DIMENSIONS. Too few specimens have been examined to permit any deductions from their dimensions, but using figures from the literature it can be seen that at a size (carapace breadth) of between 20 and 30 mm the carapace length/breadth ratio is in the range 0-47 : i to 0-49 : i. b. M. dilatatus sulcatus H. M. Edwards, 1852 Macrophthalmus sulcatusH. M. Edwards, 1852 : Alcock, 1900 ; Lenz, 1905 ; Tesch, 1915 ; Kemp, 1919 ; Chhapgar, 1957 nee Ortmann, i8g4a Macrophthalmus sandakani Rathbun, 1914 : Tesch, 1918 nee Rathbun, 1924 Macrophthalmus malaccensis Tweedie, 1937 : Crosnier, 1965 Macrophthalmus carinimanus : Lanchester, igoob MATERIAL EXAMINED. 3