LIBRARY OF THE UNIVERSITY OF ILLINOIS AT URBANA-CHAMPAIGN 590.5 FI v. 39 cop. 3 NATURAL HISTORY, SURVEY '* ' wniRAiMsrimrsmEy MAY 3UJ968 FIELDIANA . ZOOLOGY mmi Published by CHICAGO NATURAL HISTORY MUSEUM Volume 39 July 25, 1961 No. 49 On the Nomenclature of Certain Whales Philip Hershkovitz Curator, Division of Mammals The following notes were made during the course of compiling data for the section on Cetacea in a checklist of recent South Amer- ican mammals.1 Nearly all names discussed are those of cetaceans not restricted to South American waters. Generic Names for Risso's Dolphin and the Killer Whale Grampus Gray, 1828, for Risso's dolphin, and Orcinus Fitzinger, 1860, for the killer whale, are names currently recognized by most European authors. In 1933, however, Iredale and Troughton (Rec. Australian Mus., 19: 28-36) transferred Grampus Gray, 1828, to the killer whale and erected Grampidelphis for Risso's dolphin. This rearrangement was adopted by some North American mammalo- gists (cf. Miller and Kellogg, 1955, List of N. Amer. Recent Mamm., pp. 661, 662; Hall and Kelson, 1959, N. Amer. Mamm., pp. 824, 826), without heed to objections voiced by Ellerman and Morrison-Scott (1951, Checklist, Palaearctic and Indian Mamm., p. 739) and Sche- vill (1954, Jour. Mamm., 35: 124). The present conflict may be resolved by a review of the history of the technical names involved. Grampus was introduced by Gray in 1828 (Spicilegia Zoologica, 1 : 2) as a subgenus of Delphinus. It was characterized as "caput globosum; rostro depresso, longitudine capitis; dentes cylindrici, curvati." The included species were D. griseus Cuv., D. grampus, Linn, (sic), D. globiceps (= Globicephala melaena Traill), and the fol- lowing, described as new: Delphinus (Grampus) acutus (=Lageno- rhynchus acutus Gray), D. (Grampus) Heavisidii (=Cephalorhynchus heavisidii Gray) and D. (Grampus) obscurus (= Lagenorhynchus cru- ciger Quoy and Gaimard). Gray made no overt designation of a type species. In this case, the tautonym "D. Grampus, Linn [aeus?]," would automatically be- 1 Aided by a grant from the National Science Foundation. Library of Congress Catalog Card Number: 61-16636 No. 921 547 548 FIELDIANA: ZOOLOGY, VOLUME 39 come the type were it not for the fact that this name is a nomen nudum. Linnaeus never proposed it and Gray did not indicate the species of delphinid or "grampus" to which the name must be applied. The practice at the time, however, was to recognize as type of a genus either the first (or only) species mentioned or any of the species on which the generic diagnosis was based. Delphinus griseus Cuvier satisfied both requirements and was universally regarded as the type of Grampus. In 1828, Lesson (Ferussac Bull. Sci. Nat., 16: 116) reviewed Gray's article containing the description of Grampus and regarded the genus as equivalent to his own Globicephalus (sic) based on Del- phinus globiceps Cuvier (= Globicephala melaena Traill) and D. risso- anus Cuvier (= Grampus griseus Cuvier) . A second review of Gray's Spicilegia appeared the next year (Zool. Jour., 4: 497). Here the statement "typical species Delph. Grampus Linn." is made, but the anonymous reviewer failed to define the species. The name, there- fore, remained a nomen nudum. Had it been described, validation of the name would date from the anonymous author, 1829, and not from Gray, 1828. In this case, Delphinus grampus Anonymous would be a species not certainly included in the original description of Grampus and cannot be considered in determining the type of the genus (cf. Regies, Art. 30 II a). The name would also be a junior homonym of Delphinus grampus Blainville, 1817, a synonym of D. orca Linnaeus. In 1846, Gray (Zool. Voy. H.M.S. Erebus and Terror, 1, Mamm., p. 30) revised Grampus by dividing it into the four genera shown above in parentheses and leaving only Delphinus griseus Cuvier, re- named Grampus Cuvieri, in the restricted genus. This action may be construed as a selection of Delphinus griseus as type of Grampus Gray, 1828 (cf. Regies, Art. 29). Later authors, including Gray, con- ferred the tautonym "grampus" on Delphinus griseus and finally, in 1862, Reinhard (Overs. Kongl. Danske Vid. Selsk. Forh., pp. 107, 148 and footnote 1) formally declared this species the type of the genus. In 1933, Iredale and Troughton (Rev. Australian Mus., 19: 28) discovered that on page 23 of the 1922 edition of Lydekker's Guide to the British Museum Collection of Cetacea the vernacular "grampus" was mentioned once as an alternate name for the "killer," while Grampus Gray was recognized as the scientific name for Risso's dol- phin. This seemingly ambiguous state of affairs inspired Iredale and Troughton to apply prompt remedial action. The authors first averred that grampus had been used as a vernacular name for many HERSHKOVITZ: NOMENCLATURE OF WHALES 549 kinds of cetaceans but mostly, they thought, for the killer. Next, they pointed to a killer called grampus by Hunter (1787, "Phil. Trans., xvi, p. 306, ... pi. 5, figs. 1-2 [but should be 77, pt. 1, p. 373, pi. 16]"). Then followed the deduction that inasmuch "as Gray was notorious for careless writing and not reading proof carefully, it is most prob- able that the familiar 'Linn.' [in the nomen nudum Delphinus grampus Linn.] was merely a simple error for the less usual abbreviation of 'Hun.' for Hunter." Ergo, the type species of Grampus Gray, 1828, is "Delphinus grampus 'Linn.' =Hunter"= Delphinus orca Linnaeus or the killer whale. All this and more too painful to recite is sheer fantasy. Hunter never used the binomial Delphinus grampus. He used only the ver- nacular grampus for the killer and he employed the same name in the same paper for other toothed whales, including the common por- poise, Phocoena phocoena Linnaeus. Had Gray intended to base his Delphinus grampus on the grampus of Hunter, he could only have had the common porpoise in mind as this is the animal discussed under Grampus Cuvieri in his 1846 revision of the genus (Zool. Voy. H.M.S. Erebus and Terror, 1, Mamm., p. 31). As for the killer, Gray's original definition of Grampus excludes it. Indeed, Gray made the killer the type of a distinct genus for which he proposed the eminently appropriate but preoccupied name, Orca. Nomenclatorial "stability" such as was sought by Iredale and Troughton cannot be attained by electing a nomen nudum for geno- type and then defining it as fancy dictates. Grampidelphis Iredale and Troughton, 1933, proposed as a substitute for Grampus Gray, 1828, is a gratuitous synonym. Orcinus Fitzinger, 1860, never Grampus Gray, remains as the correct generic name for the killer. Still another name enters into the discussion. According to Ire- dale and Troughton (op. cit., pp. 30-31), Grayius Scott (1873, Mam- malia recent and extinct, p. 104) had already been proposed as a seemingly more appropriate generic name for Risso's dolphin. This name, however, they rejected because it was "preoccupied by Bona- parte 1856, Giinther 1858, Bate 1862." Hence, their own Grampi- delphis was offered as a substitute. I have not seen Scott's work, which is described as "an elementary treatise for the use of the Pub- lic Schools of New South Wales." If correctly cited as Grayius, the name proposed by Scott is not preoccupied by Graya Bonaparte, 1856 (a bird), Grayia Giinther, 1858 (a reptile), Grayia Bate, 1862 (a crustacean), or anything else. Thus, Grampidelphis Iredale and 550 FIELDIANA: ZOOLOGY, VOLUME 39 Troughton appears to be a superfluous as well as gratuitous synonym of both Grampus Gray and Grayius Scott. Tursiops nesarnack Lacepede, 1804, Valid Name for North Atlantic Bottlenosed Dolphin Delphinus nesarnack Lacepede (1804, Hist. nat. c£tacees, pp. xliii, 307, pi. 15, fig. 2 [animal ex Bonnaterre]) is based on the nesarnak described and figured by Bonnaterre (1789, Tabl. Encycl. M£th. Celologie, p. 21, pi. 11, fig. 1). Bonnaterre used Delphinus tursio for his nesarnak, but this name had been proposed in 1780 by Fabri- cius (Fauna Groenlandica, p. 49) for a different kind of cetacean. Bonnaterre (op. cit., p. 22) and later authors suspected as much and finally True (1903, Proc. Acad. Nat. Sci. Philadelphia, p. 313) re- jected the name completely. Delphinus nesarnack Lacepede stands, therefore, as the first valid name for the common North Atlantic bottlenosed dolphin of Bonnaterre and authors. Delphinus truncatus Montagu (1821, Mem. Wernerian Soc. Nat. Hist., 3: 75, pi. 3, skull) is a synonym. The type of D. nesarnack Lacepede is the figured animal, the model for which, according to Bonnaterre, is, or was, preserved as a mounted specimen in the Veterinary School of Alford [Maisons Alfort], near Charenton, a suburb of Paris. The type locality given by Lacepede is simply the North Atlantic but the type may have been stranded on a French coast or taken offshore. The possibility that Delphinus siculus Rafinesque, 1810, might also be a bottlenosed dolphin was considered by True (supra cit.) but its description seemed to be inadequate for positive identification. In any case this name is antedated by D. nesarnack Lacepede. The status of Delphinus carbonarius Wagner, 1846, and D. fuscus Reichenbach, 1846 Delphinus carbonarius Wagner, 1846 (Schreber's Saugth., 7: 305, pi. 352, fig. 1 [animal]) is based on the account of the "Black Fish of South Sea Whalers," by Bennett (1840, Narrative of a whaling voyage round the globe from the years 1833 to 1836, 2: 233, fig.). The same account and a skull served for the description of Globi- cephala macrorhyncha Gray (1846, Zool. Voy. H.M.S. Erebus and Terror, 1, Mamm., p. 33). The type skull, from the "south seas," was presented to the Royal College of Surgeons by Bennett but is now in the British Museum (B.M. 1946.8.9.2.; cf. Fraser, 1950, HERSHKOVITZ: NOMENCLATURE OF WHALES 551 Atlantide Report, no. 1, p. 50). Bennett's blackfish was observed on the cruise of the Tuscan "in many parallels of latitude, between the Equator and 50° N. and 35° S., in the central parts of the At- lantic and Pacific Oceans; as well as off the coast of California, and in the Indian Archipelago." It is described as wholly black, with short pectorals and massive head and with l^f teeth. Gray gives the dental formula of the type skull of macrorhyncha as | and Fraser (supra cit., p. 55) determines it as right ^, left 7j^. Delphinus fuscus Reichenbach, 1846 (Vollst. Naturg. des In- und Auslandes, Cetaceen, p. 77) is based on a uniformly blackish brown blackfish observed in the southwest Pacific by Lesson (1827, Voy. autour du monde, "La Coquille," Zool., p. 185). It is practically certain that this cetacean is also representative of the widely dis- tributed short-finned blackfish (Globicephala macrorhyncha Gray). The nomen nudum Globicephalus fuscus Hamilton (1837, in Jardine, Nat. Library, Mamm., 6: 220) is presumably another reference to the animal described by Reichenbach. All described forms in question were named in 1846 but no infor- mation is at hand as to which of the three names proposed has pri- ority. For present purposes, however, fuscus and carbonarius are arbitrarily regarded as junior synonyms of Globicephala macrorhyn- cha Gray. Before final disposition of the nomenclatorial problem is made, a fourth name must also be taken into account. Globicephala sie- boldii Gray, also described in 1846 (supra cit., p. 32), has page pri- ority over G. macrorhyncha. The name is based on the figure and skeleton of an immature blackfish first noticed under the name Del- phinus globiceps Cuvier, by Temminck (1841, Fauna japonica, p. 17, pi. 27). Fraser {in Ellerman and Morrison-Scott, 1951, Check- list, Palaearctic and Indian Mamm., p. 741) regards the skull, as indicated by the premaxillae, more like that of G. macrorhyncha or G. scammoni than that of G. melaena. Source Reference for Phocoena G. Cuvier, 1817 Authors of current checklists suggest that Phocoena Cuvier, 1817, Regne animal, 1: 279 (sometimes misspelled "Phocoena" or incor- rectly dated 1816), is unavailable but that Phocoena Cuvier, 1817, Nouveau dictionnaire d'histoire naturelle, 9: 163, is valid. Authority for the judgment "unavailable" is said to rest with Sherborn. Ex- amination of the two works cited shows the name "Phocoena Cuv." 552 FIELDIANA: ZOOLOGY, VOLUME 39 introduced in precisely the same way in both. The diagnoses and descriptions are adequate in both publications but shorter in the Regne animal. In citing Phocoena Cuvier from the Regne animal, authority for the technical name is unequivocal and there is no question of its validity under the Regies. On the other hand, authorship of the article dauphin in the Dictionnaire where the name Phocoena appears, is not always clear. Desmarest is the ostensible author but large portions were contributed by and credited to Blainville. There is no internal evidence that G. Cuvier is responsible for any part of the article. Meomeris Gray, 1847, Generic Name for Black Finless Porpoise Neomeris Gray, 1846, with type Delphinus phocaenoides Cuvier, was said to be preoccupied by Neomeris Lamouroux, 1816, described as a polyp. The conflict in names was pointed out by Palmer (1899, Proc. Biol. Soc. Washington, 13: 23), who proposed Neophocaena as a substitute. However, Thomas (1922, Ann. Mag. Nat. Hist., (9), 9: 676) showed that Meomeris Gray (1847, List osteological speci- mens, Brit. Mus., p. 36), a misprint for Neomeris, is valid as a tech- nical name and is the first available substitute for Neomeris Gray, 1846. Again, in 1925, Thomas (Ann. Mag. Nat. Hist., (9), 16: 655) called attention to an opinion that the senior homonym Neomeris Lamouroux was an alga and, as a botanical term, does not affect the status of Neomeris Gray. Unfortunately, this revival of Neomeris Gray, 1846, cannot be sustained. The same name had already been used in 1844 by 0. G. Costa (Ann. Acad. Aspiranti Nat., Naples, 2 : 81) for a genus of Annelida. It seems necessary, therefore, to rein- state the accidental Meomeris Gray, 1847, with type, Meomeris pho- caenoides (= Delphinus phocaenoides Cuvier), for the black finless porpoise. Meomeris has had scant currency but the best anatomical account of the genus with a historical summary is given under this name by G. M. Allen (1923, Bull. Mus. Comp. Zool., 65: 233-256, pis. 1-3). This author (op. cit., p. 239) cites another misprint, Phacecana, which must be added to the generic synonymy of Meomeris. The combination Phacecana [sic] phocsenoides for Delphinapterus phocae- noides Cuvier, was published in 1918 by Robinson and Kloss (Jour. Fed. Malay States Mus., 8: 79) and is duly recorded by G. M. Allen in his synonymy of Meomeris phocaenoides G. Cuvier. A third syn- HERSHKOVITZ: NOMENCLATURE OF WHALES 553 onym born of a typographical error is Nomeris Coues (1890, Century Dictionary, 4: 4449 [not seen]; 1911, 7: 4449). It was described as a subgenus of Phocaena (sic) with type P. melus (sic) (=Delphinus melas Temminck=D. phocaenoides G. Cuvier). Status of Sotalia Gray Sotalia was erected in March, 1866, by Gray (Cat. seals and whales Brit. Mus., p. 401) with type, by monotypy, Delphinus guia- nensis P. J. Van Ben&Ien. In September of the same year, Gray (Proc. Zool. Soc. London, 1866: 213) proposed Sousa as a subgenus of Steno, to contain Steno capensis Gray (a Stenella) and S. lentigi- nosus Gray, type by subsequent designation (Iredale and Troughton, 1934, Mem. Australian Mus., 6: 67). Gray (op. cit.) also proposed the subgenus Tucuxa for Steno tucuxi Gray. Authors have generally referred the types of Sousa and Tucuxa and the related species brasiliensis E. Van B£n£den, borneensis Lydek- ker, chinensis Osbeck (see below), fluviatilis Gervais and d'Orbigny (= tucuxi), plumbea G. Cuvier, and teuszii Kukenthal to the prior genus Sotalia. However, Iredale and Troughton (supra cit.) adopted Sousa for lentiginosa (and Steno gadamu Gray, a Tursiops) without mention of Sotalia. They were followed by Fraser and Purves (1960, Bull. Brit. Mus. Nat. Hist., 7, fig. 26, pi. 25 and text), who use Sousa to the exclusion of Sotalia. These authors give no clue to why or how Sousa may replace Sotalia in part or in whole. In the absence of evidence to the contrary, the name Sotalia is used here for all types and other species listed above except as noted. Sotalia chinensis Osbeck, 1765, for the Chinese White Dolphin Delphinus chinensis was proposed by ship's Chaplain Peter Os- beck in his journal of a voyage to China and the East Indies. This work, in Swedish, published in 1757, included accounts of many of the important plants and animals Osbeck saw at sea and in the coun- tries he visited. It was natural for Osbeck, a student of Linnaeus, to name and classify the species according to the master's system. Un- fortunately, Osbeck's binomials appeared one year before our start- ing-point of zoological nomenclature and the Law of Priority. There is, however, a German edition published in 1765 in Rostock. It was translated from the original Swedish by J. Gottlieb Georgi under the direction of Daniel Schreber. This translation was revised by Os- 554 FIELDIANA: ZOOLOGY, VOLUME 39 beck himself and some additions were made. The technical name for the white Chinese dolphin dated from this work is valid and may be cited as Delphinus chinensis Osbeck (1765, Reise nach Ostindien und China, p. 337). The entire original description is: "Schneeweisse Tummler (Delphinus chinensis) hupften um das Schiff, vom weiten aber konnte man sie von der gemeinen Art [i.e., Delphinus delphis] an nichts, als der weissen Farbe unterscheiden." The type was observed in the Canton River, China, November 27, 1751. The description, though brief, is adequate and the comparison apt. The white dolphin of the Chinese coast can be nothing else than Delphinus chinensis Osbeck, currently referred to the genus Sotalia. A third edition in English was translated from the German by John Reinhold Forster and published in London in 1771. The work, entitled A voyage to China and East Indies, is in two volumes. The binomial Delphinus chinensis with full description appears on page 27 of the second volume. The name Delphinus sinensis Desmarest, 1822 (Mammalogie, p. 514), adopted by Cuvier, 1836 (Hist. nat. c£tac£s, p. 213) is merely an emendation of D. chinensis Osbeck. Status of Pontoporia Gray, 1846 Pontoporia Gray, 1846, is the earliest valid generic name for the La Plata dolphin, Delphinus blainvillei Gervais. This widely circu- lated name is not a homonym of Pontoporeia Kroyer, 1842, a crusta- cean, nor has it been proven that the name is antedated by Ponto- poria Agassiz, 1846 (Nomencl. Zool., p. 305), an invalid emendation of Pontoporeia Kroyer. Stenodelphis d'Orbigny and Gervais, 1847, used by some authors for Pontoporia Gray, is a junior objective synonym. Generic Names for the Gangetic Dolphin With a Note on Sectional Names Susu Lesson, 1828 (Compl. oeuvr. Buffon, pi. 3, fig. 3 [caption], pp. 215, 440 [index]), with type, by monotypy, Susu platanista Lesson (= Delphinus gangeticus Lebeck), appears to be the earliest valid generic name for the Gangetic dolphin or susu. A second name, Platanista, was proposed in 1830 by Wagler (Nat. Syst. Amphibien, p. 35) as a classical substitute for the barbaric Susu. Wagler desig- nated Delphinus gangeticus Lebeck as type with Susa [sic] Planta- nista [sic] Lesson, a synonym. In 1825, G. Cuvier (Recherches sur les ossemens fossiles, ed. 3, pt. 1, p. 79) referred to the susu as "prob- HERSHKOVITZ: NOMENCLATURE OF WHALES 555 ablement le platanista de Pline [Hist. Nat., lib. IX, cap. XV.]" Obvi- ously, Cuvier was not coining a technical term, but Neave (1940, Nomen. Zool., 3: 780) lists it as a generic name, though with some doubt. Sousou Cuvier, 1836 (Hist. nat. c£tac£s, p. 257) and Soosoo Ham- ilton 1837 (in Jardine, Nat. Library, Mamm., 6: 254) are emenda- tions of Susu Lesson. Sousa Gray, 1866, is a synonym of Sotalia Gray, 1846 (see above, p. 553). Still another contribution to the generic synonymy of the susu is Platanistina, regarded by Neave (loc. cit.) as an "emend [ation] pro \Platan)ista Wagler 1830 [by] Gray (1846), Zool. Ereb. Terr. (Cetac), 45." In Gray's opus (Zool. Voy. H.M.S. Erebus and Terror, 1, Mamm.) Platanistina appears on page 45 as a supergeneric taxon with Platanista Wagler the only included genus. In his classifica- tion (p. 24) Gray divides the family Delphinidae "into sections by the forms of the skull, and these into genera by the form and de- scription of the teeth, and by the absence or presence of the dorsal fin." The "sections," each with its obvious type genus, are Hypero- odontina, Platanistina, Delphinina, Orcadina [from Orca, now Orci- nus], Monocerina and Iniina. These sectional or tribal names are regarded in zoological nomenclature as co-ordinate with family names. Platanistina, if recognized as a generic name, would date from Neave, 1940, who treats it as such, and not from Gray, 1846, who does not. In connection with the use of the prior name Susu, the super- generic names Susuoidea, Susuidae, and Susuinae would replace Platanistoidea, Platanistidae and Platanistinae, respectively. The Le Havre Whale and Generic Names for Beaked Whales A beaked whale stranded September 9, 1825, on the beach of Sainte-Adresse near Le H&vre, was regarded by Blainville (1825, Bull. Soc. Phil., Paris, 4: 139) as representative of the "bottle-head" of Dale (1732, History and antiquities of Harwich and Dovencourt, ed. 2, p. 411, fig. 14). A second notice by Blainville (1826, op. cit., 5: 193, pi. for August, 2 figs.) included a figure of the animal and another of its skull. The same year, F. Cuvier (1826, in Geoffroy and Cuvier, Hist. Nat. Mamm., 6, livr. 53, pi.) published a descrip- tion and colored figure of the Le Havre whale under the name dau- phin de Dale. Blainville's figure resembles the engraving of Dale's "bottle-head" or bottlenosed whale (= Hyperoodon ampuUatus Forster, 556 FIELDIANA: ZOOLOGY, VOLUME 39 1770) while Cuvier's representation is that of a beaked whale (Meso- plodon of authors) . The skull, however, as figured and described by Blainville and later authorities (cf. Gervais, 1859, Zool. Pal. Fran- caises, ed. 2, p. 291, pi. 40, fig. 1) appears to be conspecific with that of the beaked whale, Physeter bidens Sowerby, described in 1804. The first technical name for the Le Havre beaked whale, Heterodon dalei, proposed by Lesson in 1827 (Man. Mamm., p. 419) perpetuated the erroneous association with Dale's "bottle-head." Lesson fur- ther aggravated the misrepresentation by including Delphinus edent- ulus Schreber (= Hyperoodon ampullatus Forster) in the synonymy.1 The following year, Lesson (1828, Compl. oeuvr. Buffon, 1, C£ta- c£s, pp. 149, 155, 440, pi. 3, fig. 1) erected Aodon to contain Heterodon dalei, now Aodon dalei (op. cit., pi. 3, fig. 1) with synonyms (p. 155) "Delphinus edentulus, Schreb., Desm.; dauphin de Dale, de Blain- ville. F. Cuvier." The entire description is based on the Le Havre whale, and Aodon Lesson, 1828, would be the prior generic name for beaked whales were it not preoccupied by Aodon Lacepede, 1798, a genus of fish. Heterodon Blainville (1817, in Desmarest, Nouv. diet, d'hist. nat., 9: 175) used by Lesson in the original description of dalei, is a composite of Monodon Linnaeus, Hyperoodon Lacepede and the un- identifiable Epiodon Rafinesque (see below, p. 564). Its type, Del- phinus butskode Blainville (now selected ;=D. butskopf Bonnaterre, 1789) is a Hyperoodon and the generic name itself is invalidated by Heterodon Latreille, 1801, a reptile. The next name for consideration is Nodus Wagler (1830, Nat. Syst. Amphibien, p. 34, and footnotes 2, 3). It was explicitly pro- posed as a substitute for the preoccupied Aodon Lesson. Its type species, identical with that of Aodon, is cited thus: "Delphinus edent- ulus Schreb., Cuv. Mammif. Liv. 53. (Heterodon Dalei Less. Man. de Mammal, p. 419. Id. Aodon Dalei Oeuvr. de Buff. p. 155. t. 3. f. 1)." Wagler's concept of Delphinus edentulus in the place cited is clearly not that of Schreber, which is a bottlenosed whale, but that of Lesson and Cuvier, which is the beaked whale of Le Havre. Inas- much as Nodus Wagler is erroneously listed as a synonym of Hyperoo- don by modern authors (cf. Ellerman and Morrison-Scott, 1951, ^Herman and Morrison-Scott (1951, Checklist, Palaearctic and Indian Mamm., p. 723) list the Le Havre whale with the original name Heterodon dalei in the synonymy of Hyperoodon ampullatus Forster, and assign it the erroneous type locality, "Harwich, England." On page 725 (op. cit.) the same animal, under the name Aodon dalei Lesson, is correctly listed in the synonymy of Mesoplodon [ = Nodus] bidens Sowerby but the type locality given is "North European waters." HERSHKOVITZ: NOMENCLATURE OF WHALES 557 Checklist, Palaearctic and Indian Mamm., p. 722) it cannot be over-emphasized that the true Delphinus edentulus Schreber is not the type of the genus. It is only a name wrongly applied to the Le Havre beaked whale, Aodon dalei Lesson (=Heterodon dalei Lesson) which is the type. For all the names and authors involved in the cited synonymies, no other species is included in either Aodon or Nodus as originally proposed and no other species can be designated type. Nodus Wagler, therefore, appears to be the earliest valid name for beaked whales currently referred to Mesoplodon. In his remarks on Aodon Lesson, Wagler (op. cit., p. 34, footnote 3) used the misspelling or typographical error, Anodon. As defined, Anodon Wagler (not Aodon Lesson) includes only the true Delphinus edentulus Schreber, here designated type, and Delphinus butzkopf [sic] Bonnaterre. Anodon, thus, is a synonym of Hyperoodon Lacepede. It is also a junior homonym of Anodon Smith, 1829, a reptile. The next generic name for the Le Havre whale is Micropterus Wagner, 1846 (Schreber's Saugth., 7: 281, 352). Its type, Delphinus micropterus G. Cuvier (1829, Reg. anim., ed. 2, 1: 288), is merely a new name for Heterodon dalei Lesson. Micropterus Wagner is in- validated by Micropterus Lacepede, 1802, a genus of fish. In 1849, however, Eschricht (Kongl. Danske Vidensk. Selsk. Skrift., (5), 1 : 97) independently erected the same generic tautonym for Delphinus mi- cropterus Cuvier but wrote it Micropteron. This form of the name is valid. In 1850, Gervais (Ann. Sci. Nat., Paris, (3), 14: 16) proposed two generic names for beaked whales. The first, Dioplodon, with type by original designation, Delphinus densirostris Blainville, appears on lines 19-20 of the page cited. The second, Mesoplodon, with type by original designation, Delphinus sowerbensis Blainville (=Physeter bidens Sowerby) is on line 26 of the same page. Subsequently, at least six more generic names were proposed for beaked whales by various authors. They are included in the following synonymy. Genus NODUS Wagler Aodon Lesson, 1828, Compl. oeuvr. Buffon, 1, C6taces, pp. 155, 440, pi. 3, fig. 1 (animal with caption) — type species, Aodon Dalei Lesson [ = Physeter bidens Sowerby], by monotypy; generic name preoccupied by Aodon Lacepede, 1798, a genus of fish. Nodus Wagler, 1830, Nat. Syst. Amphibien, p. 34 — new name for Aodon Lesson, preoccupied; type species, "Delphinus edentulus Schreb., Cuv. Mammif. Livr. 53. (Heterodon Dalei Less. Man. de Mammal, p. 419. Id. Aodon Dalei Oeuvr. deBuff. p. 155. t. 3. f. 1)" [ = Physeter bidens Sowerby]. 558 FIELDIANA: ZOOLOGY, VOLUME 39 Micropterus Wagner, 1846, Schreber's Saugth., 7: 281, 352 — subgenus of Delphinus; type species, D. micropterus Cuvier [ = Physeter bidens Sowerby] by tautonymy and monotypy; generic name preoccupied by Micropterus Lacepede, 1802, a genus of fish. Micropteron Eschricht, 1849, Kongl. Danske Vidensk. Selsk. Skrift., (5), 1 : 97 — type species, Delphinus micropterus Cuvier [=Physeter bidens Sower- by], by tautonymy and monotypy. Dioplodon Gervais, 1850, Ann. Sci. Nat., Paris, (3), 14: 16 (lines 19-20)— type species, Delphinus densirostris Blainville by original designation and mon- otypy; Gervais, 1850, Compt. rend. Acad. Sci. Paris, 31: 512 (extract of previous article). Mesoplodon Gervais, 1850, Ann. Sci. Nat., Paris, (3), 14: 16 (line 26) — type species, Delphinus sowerbensis Blainville [ = Physeter bidens Sowerby] by original designation and monotypy. Mesodiodon Duvernoy, 1851, Ann. Sci. Nat., Paris, (3), Zool., 15: 41— included species, Mesodiodon Sowerbyi Duvernoy [type= Delphinus sowerbensis Blain- ville= Physeter bidens Sowerby], Mesodiodon micropterum [ = Delphinus mi- cropterus Cuvier = Physeter bidens Sowerby], Mesodiodon densirostre ^Del- phinus densirostris Blainville], Mesodiodon longirostre, "qui est le Ziphices [sic] longirostris de Cuvier [fossil]." Mesiodon Gray, 1866, Cat. seals and whales Brit. Mus., p. 349 — misspelling of Mesodiodon Duvernoy, 1851. Diplodon Gray, 1866, Cat. seals and whales Brit. Mus., p. 349 — misspelling of Dioplodon Gervais, 1850. Dolichodon Gray, 1866, Cat. seals and whales Brit. Mus., p. 353 — subgenus of Ziphius Cuvier; type species, Ziphius layardii Gray, by monotypy. Callidon Gray, 1871, Ann. Mag. Nat. Hist., (4), 7: 368 — type species, Meso- plodon Giintheri Krefft [= Ziphius layardii Gray], by monotypy. Neoziphius Gray, 1871, Suppl. Cat. seals and whales Brit. Mus., p. 101 — type species, Neoziphius europaeus Gervais [= Dioplodon europaeus Gervais], by monotypy. Oulodon von Haast, 1876, Proc. Zool. Soc. London, 1876: 457 — type species, Mesoplodon grayi Haast, by monotypy; Von Haast, 1877, Trans. New Zea- land Inst., 9:450 (characters). Paikea Oliver, 1922, Proc. Zool. Soc. London, 1922: 574 — included species, Berardius hectori Gray [type, by original designation], Mesoplodon mirus True. Status of Dioplodon europaeus Gervais Dioplodon europaeus Gervais, 1852 (Zool. Pal. Fr., ed. 1 [1848- 1852], p. 4) is said to be a nomen nudum (cf. Ellerman and Morrison- Scott, 1951, Checklist, Palaearctic and Indian Mamm., p. 725). ' I 1 Cited here as "1852 Dioplodon europaeus Gervais. Zool. Pal. Fr. 2, text to pi. 40, nom. nud." If correctly cited, it is puzzling that a binomial proposed with text to an illustration would be a nomen nudum. Is it possible that the "plate 40" is the same referred to by True, (1907, cited below)? HERSHKOVITZ: NOMENCLATURE OF WHALES 559 have not seen the first cited work. The same name, however, appears on pages 289, 298 and 299 of the second revised edition of Gervais' work, published in 1859. A history of the type specimen with some allusions to its characters is given here but nothing else which can be construed as a valid description. It seems, therefore, that neither the first nor the second edition of the Zoologie et Paleontologie Fran- caises can be cited for the original description of Dioplodon europaeus Gervais. The same opinion was expressed in 1907 by True (Science, 26: 796). In 1910, however, True reverted to the name Mesoplodon europaeus in his account of the ziphiids in the collection of the U. S. National Museum (Bull. U. S. Nat. Mus., 73: 11). No explanation was given, but he cited the two editions of Gervais' work as authority for the name and added "pi. 40, figs. 3-6," to the 1859 reference. The figures in question, in the Atlas to the second edition of Zoologie et PaUontologie Francaises, are of the skull and mandible of an Indian Ocean representative of a correctly identified and captioned Dioplo- don densirostris Blainville! Notwithstanding the disputed bibliographic references, the name Dioplodon europaeus was published with a valid description in 1855, by Gervais in the second volume (p. 320) of his Histoire naturelle des mammifkres (Paris). In 1863, Van B£n£den (Mem. couronnes et autres Mem., Acad. Roy. Belgique, 16: 18) also described Dioplodon europaeus in a comparison of the type skull with his Ziphius indicus. The name for the European beaked whale stands, therefore, as Nodus europaeus Gervais, 1855, with Dioplodon gervaisi Deslongchamps, 1866, a synonym. Type Species of Balaenoptera Lacepede The generic name Balaenoptera, as conceived by Lacepede (1804, Hist. nat. c£tacees, pp. xxxvi, 114), consists of two subgenera and four species. The first, or premier sous-genre, comprises only Balaenoptera gibbar [=Balaena physalus Linnaeus]. The second sous-genre includes Balaenoptera jubartes Lacepede [=Balaena musculus Linnaeus], Bal- aenoptera rorqual Lacepede [=Balaena physalus Linnaeus], and Balae- noptera acutorostrata Lacepede. These species are numbered 1 to 4, respectively. In the Sonnini edition of Lacepede's work, also pub- lished in 1804, the phrase premibre espbce is used to indicate the type of each genus. Other species of the same genus are merely numbered consecutively as in the original edition. The premikre esp&ce, or type, of the first or typical section of the genus Balaenoptera is the fin whale, the "baleinoptere [or] baleine a nageoires," Balaenoptera gibbar 560 FIELDIANA: ZOOLOGY, VOLUME 39 Lacepede, by virtual tautonymy as well as monotypy. This is the sense given by Lacepede himself (op. cit., footnote, p. xxxvi; Sonnini ed., footnote, p. 193). Balaenoptera rostrata Fabricius (not Miiller) "considered" by Gray (1847, Proc. Zool. Soc. London, 1847: 90) as type of the genus is a subjective synonym of Balaenoptera acutorostrata Lacepede. As such, it can be regarded as the subsequently designated type of only the second sous-genre of Balaenoptera. Flower (1865, Proc. Zool. Soc. London, 1864: 395) followed Gray by clearly designating B. rostrata Fabricius as the type of "Balaenoptera Lacep. pars." The following year Gray (1866, Cat. seals and whales Brit. Mus., p. 382) raised this "pars" of Balaenoptera Lacepede, typified by Balaenoptera ros- trata Fabricius (or B. acutorostrata Lacepede), to full generic rank and named it Fabricia (not Fabricia Blainville, 1828, a genus of Vermes) . Balaenoptera gibbar Lacepede (Balaena physalus Linnaeus) remains, as it always has been, the first designated type of the typical section of Balaenoptera Lacepede. Status of Agaphelus Cope Agaphelus Cope is currently listed as a synonym of Eschrichtius Gray. This appears to be incorrect. In 1868 Cope (Proc. Acad. Nat. Sci. Philadelphia, 20: 159) ex- amined "a portion of a specimen of [he thought] the Scrag Whale of Dudley, Balaena gibbosa of Erxleben, and ascertained that it rep- resented a genus not previously known." Cope named the genus Agaphelus. He then appended a second species, Agaphelus glaucus Cope, the "gray whale of the coasts of California." In 1869, Cope (op. cit., 21 : 15) decided that Agaphelus glaucus was generically different and made it the type of a new genus, Rhachianectes Cope. Thus, Agaphelus was left with its originally designated type, the "portion of a specimen" identified with "Balaena gibbosa Erxleben," or Agaphelus gibbosus Cope. It was not until 1884 that Cope (Amer. Nat., 18: 1124) revealed that the bones he identified as Balaena gibbosa "are probably those of Balaenoptera rostrata [=B. acutoro- strata Lacepede]." If this determination is final, then Agaphelus Cope is an objective synonym of Fabricia Gray (see above) and, like it, falls into the synonymy of Balaenoptera Lacepede. The history of the scrag whale and the species assigned to Aga- phelus were reviewed by Van Diense and Junge (1937, Temminckia, 2: 178). Evidently, these authors (op. cit., p. 181) concluded that since the type species of Agaphelus proved to be a Balaenoptera, the HERSHKOVITZ: NOMENCLATURE OF WHALES 561 next species, Agaphelus glaucus, moved into the void and became the new type of the genus. The Regies provide, however, that a genus stands or falls on the basis of its original type species and that the type species cannot be altered or replaced. Agaphelus glaucus Cope, in spite of its name and the nomenclatorial vicissitudes of Agaphelus gibbosus Cope (not Erxleben) was not and never can be, type of Agaphelus. For an unnecessarily drastic recommendation for dis- position of Agaphelus Cope as a synonym of Balaenoptera see True (1904, Smithsonian Contr., 33: 105). Balaena glacialis Miiller, 1776, for North Atlantic Right Whale Balaena glacialis Miiller, 1776 (Zoologiae Danicae Prodromus, seu animalium danicae et norvegiae indigenarum. . . , p. 7), appears to be the earliest name for the North Atlantic right whale. The same name is cited in current lists but dated from Borowski, 1781, and, in older works, from Bonnaterre, 1789. Muller's description of the right whale consists of the following indications: "B. glacialis. Norwegian] Sild-Qval, Lille-Hval, Nord- Kaper. Eg. Gr. c. 6 [=Hans Egede, 1742, Des alten Gronlandes Naturell-Historie, chap. 6], Aph. 3, 594 [=H. von Aphelen, Bomare's Natur Historic vol. 3, p. 594]." These indications are followed by an asterisk(*), meaning that the species is not described in a Linnaean work and a dagger(f), meaning that the animal is mentioned by other authors. I have not seen the basic references cited by Miiller for the account of B. glacialis. There can be no doubt, however, regarding the applicability of this name to the nordcaper of Norwegians, i.e., the right whale of North Cape, Norway. The only other similar species, the Greenland right whale, Balaena mysticetus, is listed as a distinct species by Miiller on page 6 of his "Prodromus." Balaena japonica Lacepede, 1818, for North Pacific Right Whale Balaena japonica Lacepede, 1818 (Mem. mus. d'hist. nat., Paris, 4: 469, 472), from Japanese waters is listed as a dubious synonym of Eubalaena sieboldii Gray, 1864, by Ellerman and Morrison-Scott (1951, Checklist, Palaearctic and Indian Mamm., p. 718) and by Hall and Kelson (1959, Mamm. N. Amer., p. 840). There is no need for ambiguity here. If B. japonica can be identified at all as a 562 FIELDIANA: ZOOLOGY, VOLUME 39 Eubalaena, it must necessarily be the North Pacific representative of this genus. Actually, the original description and classification of Balaena japonica which follows, leaves no doubt on this score: "L'event est place" un peu au-devant des yeux; la nageoire caudale est grande; on voit sur le museau trois bosses garnies de tuberositas, et placees longitudinal ement; la couleur generate est noire; le ventre est d'un blanc eclatant, et cette grande place blanche est comme festonnee profondement dans son contour; les machoires, les bras ou nageoires pectorales et la caudale sont bord£s de blanc; des lignes courbes, noires et tres-fines relevent le blanc qui est autour des yeux et de la base des pectorales; on distingue des groupes de petites taches blanches sur la machoire inferieure; et d'autres petites taches de la meme couleur sont repandues sur le museau." The taxonomic position of B. japonica is shown in a key with diagnoses (op. cit., pp. 472-473) of the genera, subgenera and species of baleen whales. This key is proposed as a supplement to the general classification, or Tableau, in Lacepede's classic Histoire naturelle des cetacees, published in 1804. Balaena japonica is here included among the right whales. Its systematic position is given as "apres la Baleine nord caper [= Eubalaena glacialis]," which, in the original Tableau, follows the premiere espece, or type, Balaena mysticetus. This is precisely the arrangement used in modern check- lists. The only possible objection to recognition of Balaena japonica Lacepede as the earliest valid name for the North Pacific right whale may be the fact that its description is based on a drawing by a Japanese artist and not on the actual specimen. Such objection, if made, would be unrealistic. The original descriptions of nearly all recognized species of large whales are based on nothing better. Balaena sieboldii Gray, 1864 (Ann. Mag. Nat. Hist., 14: 349) cur- rently substituted for japonica, is itself a third hand description. Its basis is a bibliographic reference to Balaena antarctica Temminck, 1841 (Fauna Japonica, p. 18, pis. 28, 29), which, in turn, is based on drawings copied from a porcelain model made by a Japanese artist! These copies of the original figure of the real animal agree in all essential characters with japonica Lacepede. This last name for the North Pacific right whale was well established during the last cen- tury and most authors (cf. Van Ben£den and Gervais, 1880, Ost£- ographie des c£tacees, p. Ill) include sieboldii Gray in its synonymy. Valid characters for specific separation of the right whales of the North Atlantic (glacialis Miiller, 1776), the North Pacific (japon- HERSHKOVITZ: NOMENCLATURE OF WHALES 563 ica Lac£pede, 1818), and the southern hemisphere {australis Desmou- lins, 1822), have not been demonstrated. Tomilin (1957, Mamm., eastern Europe and northern Asia, 9 : 75) treats sieboldii [=japonica] as a subspecies of Eubalaena glacialis while Nishiwaki (1957, Coll. Rep. Fish. Sci., Tokyo, p. 150) recognizes the taxon E. glacialis ja- ponica Lac^pede. The same relationship to E. glacialis probably obtains in the case of australis. Balaena lunulata Lacepede, 1818, as described and classified in the same work (supra cit., pp. 470, 473) is another Japanese right whale, indistinguishable from B. japonica, which has page and line priority. The name Balaena antarctica, used by Temminck (supra cit.), was originally proposed by Lesson (1828, Compl. oeuvr. Buffon, 1, C6taces, p. 391) as a substitute for Balaena australis Desmoulins, 1822, based on the South African right whale. Status of Anarnak Lacepede, 1804, and Ancylodon Illiger, 1811 The genus Anarnak Lacepede, 1804 (Hist. nat. celacees, pp. xxxviii, 164) is based solely on Anarnak groenlandica Lacepede, which is a substitute name for Monodon spurius Fabricius, 1780 (Fauna Groen- landica, p. 31). Ancylodon Illiger, 1811 (Prodr. Syst. Mamm. Avium, p. 142) is based on the same species. Current checklists (cf. Eller- man and Morrison-Scott, 1951, Checklist, Palaearctic and Indian Mamm., p. 722; Hall and Kelson, 1959, Mamm. N. Amer., p. 811) list Anarnak as a doubtful synonym of Hyperoodon Lacepede and its type as a species inquirenda. On the other hand, Ancylodon is cited as a clear synonym of Hyperoodon and its type species is equated with Balaena ampullata Forster, 1770. Monodon spurius Fabricius (= Anarnak groenlandica Lacepede) is characterized by a dorsal fin, black, elongate body and two small teeth in the upper jaw. Were it not for the probable lapsus calami of locating the teeth in the upper instead of the lower jaw, identity of Monodon spurius with Balaena ampullata would be plausible and page priority of Anarnak Lacepede, 1804, over Hyperoodon Lace- pede, 1804, indicated. In view of the discrepancy, however, it is best to conserve the currently recognized Hyperoodon and treat Anarnak Lacepede, Ancylodon Illiger and their common type species, Mono- don spurius Fabricius, as Incertae sedis. Anarnacus Dumeril, 1806 (Zool. analytique, p. 28) is merely an emendation of Anarnak Lacepede and follows it as an objective syn- 564 FIELDIANA: ZOOLOGY, VOLUME 39 onym. Other emendations proposed by different authors in various works are Anarnac and Anarnakus. Status of Delphinorhynchus Blainville, 1817, and Rhamphocetus Gloger, 1842 Delphinorhynchus Blainville, 1817 (in Desmarest, Nouv. diet, d'hist. nat., Paris, 9: 151) was proposed as a subgenus of Delphinus. The originally included species are D. geoffrensis Blainville (=Inia geoffrensis Blainville), D. shawensis Blainville (= Delphinus gangetica ~Leheck= Susu gangetica Lebeck), D. pernettensis Blainville (=Sten- ella pernettensis Blainville), and D. coronatus Freminville (1812, Bull. Soc. Phil. Paris, 3: 71), now designated type of the genus. Delphinus coronatus Freminville is said to be 30-36 feet long and 15 feet in girth, its head rounded, with two yellow concentric circles on top, beak pointed, with 48 small, sharp, conical teeth in the lower jaw, 30 in the upper. It inhabits the Arctic seas from 74° northward and is particularly abundant in waters around Spitzbergen. Certain characters and the geographic distribution of Delphinus coronatus have led authors to include the species, albeit with a query, in the synonymy of Hyperoodon ampullatus Forster. This allocation would imply that Delphinorhynchus should also be listed as a dubious synonym of Hyperoodon. It is obvious, however, that D. coronatus is unidentifiable and possibly mythical. Confusion and the false im- plication of pending enlightenment are avoided by consigning Del- phinorhynchus Blainville with its type, Delphinus coronatus Fremin- ville, to the limbo of Incertae sedis. Rhamphocetus Gloger, 1842 (Hand- und Hilfsbuch der Naturge- schichte, 1: xxxiv, 169), with type, by monotypy, R. coronatus (= Delphinus coronatus Freminville), is an objective synonym of Del- phinorhynchus Blainville. Status of Epiodon urganantus Rafinesque, 1814 The type of Epiodon urganantus Rafinesque (1814, Precis des de- couvertes et travaux somiologiques . . ., p. 13) was sighted in the Mediterranean Sea near Sicily. It is said that it lacked a dorsal fin, that it had several upper but no lower teeth and that its upper jaw extended forward beyond the lower. Nothing like it has been seen before or since. The generic and specific names Epiodon and E. urga- nantus are currently classified as dubious synonyms of Ziphius G. HERSHKOVITZ: NOMENCLATURE OF WHALES 565 Cuvier and Z. cavirostris G. Cuvier, respectively (cf. Ellerman and Morrison-Scott, 1951, Checklist, Palaearctic and Indian Mamm., pp. 723, 724). In my opinion, Epiodon urganantus Rafinesque has nothing to do with any ziphiid. The animal is unidentifiable and possibly mythical.