CHAPTER 12 Relationships of Lower Euteleostean Fishes G. DAVID JOHNSON COLIN PATTERSON National Museum of Natural History Natural History Museum Smithsonian Institution London, England Washington, D.C.- We all make mistakes; then we're sorry. What are the relationships of and within the Os- Popular song meroidei? (6) What are the relationships of and within Salmonidae? (7) Where does Lepidogalaxias belong? (8) What are the relationships within stomiiform fishes? (9) What of the Myctophoidei, as recognized by I. Introduction Greenwood et al. (1966, i.e., Aulopiformes and Myc- tophiformes in current terminology)? In that agenda, In the first Interrelationships of Fishes lower eutel- items (8) and (9) are treated elsewhere in this volume eosts, or "protacanthopterygians" as they were then and do not concern us, but items (1) through (7) do. called, were omitted, with only a comment in the Some classifications and/or cladograms of lower eu- Preface citing Weitzman (1967, on osmeroids and teleosts, dating back to the first application of cladistic stomiatoids), McDowall (1969, on osmeroids and ga- method, are summarized in Fig. 1. As is obvious from laxioids), Rosen and Greenwood (1970, on gonoryn- incongruence between all the patterns in Fig. 1, there chiforms and ostariophysans), Greenwood and Rosen has been protracted argument on how lower euteleos- (1971, on argentinoids and alepocephaloids), and Nel- tean groups are interrelated, how they are related to son (1970b, on salangids and argentinids; 1972, on neoteleosts (stomiiforms and eurypterygians, John- esocoids and galaxioids). son, 1992), and what group is basal to other euteleosts. Ten years later, in Ontogeny and Systematics of Fishes, The most substantial treatment of these problems is Fink (1984a) summarized the history of protacantho- in Begle's (1991,1992) cladistic analyses of Osmeroidei pterygians as "erosion" and "attrition, most notably (1991) and Argentinoidei (1992) (Fig. 1G). Begle's two at the hands of Rosen (1973)" [in the first Interrelation- papers resulted in the cladogram in Fig. 2, in which ships of Fishes]. Fink then saw the problems as these: the terminals are the genera or higher taxa sampled in (1) What are the relationships of the Esocoidei? his matrix. His classification (Begle, 1991, fig.l; 1992, (2) What are the relationships of the Ostariophysi? Do table 2) to family level, without ranks above the super- these fishes lie above or below the Esocoidei? (3) What family and sequenced according to the conventions is the pattern of relationships among the traditional of Wiley (1981), was as follows: "salmoniform" taxa, exclusive of Esocoidei and Ostar- iophysi? (4) What are the relationships of and within Euteleostei the Argentinoidei (sensu Greenwood and Rosen, Esocae 1971, i.e., argentinoids plus alepocephaloids)? (5) Ostariophysi sedis mutabilis Copyright © 1996 by Academic Press, Inc. INTERRELATIONSHIPS OF FISHES 251 All rights of reproduction in any form reserved. 252 G. DAVID JOHNSON AND COLIN PATTERSON =Alepocephaloids T=LepidogaIaxias 4 |=Esocoids MEsocoids L-1 ir=Argentinoids ij=Alepocephaloids L= =Galaxioids ' " Argentinoids =Osmerids j=Galaxiids HSalangids ii=J=Salmonoids =Salmonoids F=Osmerids! —iL^MSalangids n" L-RiRetropinnids B j=Esocoids || ipOstariophysi ^U i?=Argentinoids =L=Alepocephaloids l=Esocoids H=Osmeroids =j| if=Ostariophysi =4=Salangids |=Neoteleostei l!=Galaxioids ij=Salmonoids fr=Lepidogalaxias MpOsmeroids JL=Salmonoids I |[=Argentlnoids IPNeoteleosts Alepocephaloids n=Salnionoids =Esocoids i, Esocoids =j| |]=Neoteleosts 4 ir=Neoteleosts L=j| ij=Salmonoids W ij=Alepocephaloids 4 |=Alepocephaloids I, " Argentinoids i, ' Argentinoids W i|=Osmeroids W ij=Salangids L=L=Galaxioids ii " Osmerids ILJI ^Galaxioids =Lepidogalaxias -Esocoids 4 |?=Ostariophysi =ip=oaiSalmonoids Neoteleosts )stariophysi Argentinoids —=Clupeomorpha Alepocephaloids pEsocoids smerids —I r]=Neoteleosts Retropinnids W ^Alepocephaloids ^Lepidogalaxias |]=L=Argentinoids W if=Salangids L=j| ip=Salmonoids LJ=Galaxiids =U=Osmeroids FIGURE 1 Cladograms or classifications of lower euteleostean fishes summarized as branching diagrams. Common names (e.g., alepocephaloids and esocoids) are used in- stead of formal taxon names to emphasize the equivalence of groups that may be given different ranks, and so names with different terminations, in classifications. (A) G. J. Nelson (1970b). (B) Rosen (1974). (C) Fink (1984b). (D) Sanford (1990). (E) Williams (1987; summarized in J. S. Nelson, 1994, p. 175). (F) J. S. Nelson (1994). (G) Begle (1991, 1992). (H) Patterson (1994). Salmonoidei sedis mutabilis Alepocephalidae (including Bathy- (Neoteleostei + Osmerae) sedis mutabilis laconidae, Bathyprionidae, Leptochi- Neoteleostei lichthyidae, Platytroctidae) Osmerae Osmeroidei Argentinoidei Osmeroidea Argentinoidea Osmeridae (including Pleco- Argentinidae glossidae) Microstomatidae Galaxioidea Bathylagidae Retropinnidae (including Proto- Opisthoproctidae troctidae) Alepocephaloidea Lepidogalaxiidae 12. Relationships of Lower Euteleostean Fishes 253 lf=Outgroups we decided that we must check each character against =Salmouoids Esocoids specimens. At the same time, we found that our own =Neoteleosts survey of lower euteleostean intermusculars (Patter- F=AplocMton S=Galaxiidae son and Johnson, 1995) was deficient both in sampling Jjovettia and in the quality of our material, casting doubt on alangidae the pairing of salmonoids and osmeroids that we pro- Lepidogalaxias F=Pmtotroctes posed. Given this disquiet about both recent sources 4 F=Retropinna of published information, Begle's and our own, we u=J=StokeIlia 'Thaleichthys were thrown into a detailed survey of lower eutel- t=Osmenis eosts, limited only by time and by the availability of ii '' Plecoglossus 4 tf=Hypomesus specimens. On sampling, we limited ourselves princi- IW *=MalIotus pally to the 33 taxa in Begle's sample (Fig. 2), but we L=II •=AllosmeTus excluded the alepocephaloid Bathyprion (unavailable) Spirinchus w=Glossanodon and added more argentinoids (bathylagids, opistho- ^Argentina proctids, and microstomatids, which Begle did not w=Bathylagus '^Opisihoproctus sample) and more platytroctid alepocephaloids. On Leptochilichthys characters, we concentrated on the 108 used by Begle •Bathylaco Narcetes (1992), the majority of them taken from the literature Rouleina [e.g., nos. 1-28, 54, 55, from Fink (1984b); nos. 30-38, w=Alepocephalus 51, 63, from Howes and Sanford (1987b)], because we *=Bathyprion Bathytroctes were driven to discover every detail of his miscoding Leptoderma or misconduct. But during our survey almost 100 other F=Talismama L=Platytroctidae characters came to light. In presenting our data, we originally had a section FIGURE 2 Result of Begle's (1991, 1992) cladistic analyses of a criticizing our own work on the intermusculars, a sample of lower euteleosts. longer critique of Begle's 108 characters, and a section describing new characters. Referees and others pointed out that this sequence gave too much atten- Salangidae (including Sunda- tion to our critique of Begle's work and artificially salangidae) separated characters into three different sets. We have Galaxiidae (including Aplochi- therefore organized the primary data into a single tonidae) survey of characters structured by anatomical region. The original plan for this volume was that Douglas We then discuss in sequence the outgroup and in- P. Begle should write about lower euteleosts since he group relationships of osmerids [Osmeridae and had covered the ground in his two papers (Begle, Salangidae of Begle (1991)], osmeroids [= Osmerae 1991, 1992). However, in 1993 Begle left ichthyology of Begle (1991)], argentinoids and alepocephaloids to move into computer programming. To fill the gap, [= Argentinoidei of Greenwood and Rosen (1971)], we agreed to take on lower euteleosts, believing that salmonoids, esocoids, and euteleosteans as a whole. we had a new angle on them from our work on inter- We do not include the "salangid" Sundasalanx, only musculars (Patterson and Johnson, 1995). In particu- member of the Sundasalangidae (Roberts, 1984), in lar, the discovery of cartilaginous epicentrals in sal- our study because it is not euteleostean (Darrell Sie- monoids and osmeroids, together with the absence bert, 1996, and personal communication). of ossified epipleurals in both, implied that they might We dedicate this work to two absent friends, be sister-groups (Patterson and Johnson, 1995, p. 26, Humphry Greenwood (1927-1995), senior editor of fig. 9). In arguing for that proposal, we criticized the the first Interrelationships, and Donn Rosen (1929- characters supporting some of the higher-level rela- 1986), the length of whose chapter in that book was tionships in Begle's cladogram, but we anticipated justified by the information it contained. May we that his conclusions at lower levels (e.g., within os- be so excused. merids or within galaxioids) would withstand criticism. However, when we began checking Begle's character descriptions and coding, we found many errors. Be- II. Character Survey cause Begle's work resulted in the only fully resolved phylogeny of lower euteleosts, and because the char- The following survey of 200 characters is organized acters that he used included the great majority of those by anatomical region or system, beginning with the previously used in systematic studies of those fishes, braincase and ending with reproductive and soft ana- 254 G. DAVID JOHNSON AND COLIN PATTERSON tomical features. One purpose of this