Comments on Begle's "Monophyly and Relationships of Argentinoid Fishes" Author(s): Colin Patterson and G. David Johnson Source: Copeia, Vol. 1997, No. 2 (May 13, 1997), pp. 401-409 Published by: American Society of Ichthyologists and Herpetologists Stable URL: http://www.jstor.org/stable/1447761 Accessed: 24/05/2010 15:26 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/action/showPublisher?publisherCode=asih. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact support@jstor.org. American Society of Ichthyologists and Herpetologists is collaborating with JSTOR to digitize, preserve and extend access to Copeia. http://www.jstor.org SHORTER CONTRIBUTIONS Copeia, 1997(2), pp. 401-409 Comments on Begle's "Monophyly and Relationships of Argentinoid Fishes" COLIN PATTERSON AND G. DAVID JOHNSON In 1992, Begle published a paper on the in- terrelationships of argentinoid fishes, with com- ments on the relationships of other lower eu- teleosts. The study was based on a matrix of 108 characters scored in 32 taxa and a hypothetical outgroup (a row of zeros). That matrix was an expansion of one in an earlier paper (Begle, 1991) comprising 84 characters scored in 19 taxa and an outgroup. Begle's (1992) data dif- fered by including 24 additional characters, modifying a few of the previous 84, and sam- pling 14 taxa of Argentinoidei (alepocephaloids and argentinoids), a group treated as a single terminal in the 1991 paper. In a revision of the interrelationships of lower euteleosts (ohnson and Patterson, 1996), we had to pay close atten- tion to Begle's two papers and found his two matrices to be rife with errors. We have a note (Patterson and Johnson, 1997) detailing errors in Begle's 1991 matrix. Here we concentrate on his 1992 matrix, the characters newly intro- duced in it, and his conclusions. Appendixes 1 and 2 reproduce Begle's (1992) matrix, togeth- er with our corrected coding and a summary of differences between the two versions. A detailed commentary is published elsewhere (ohnson and Patterson, 1996), and there is an abbrevi- ated commentary on some characters in Appen- dix 2 and the text below. Results.-When we ran Begle's (1992) matrix, as published, on Hennig86, version 1.5 (. S. Far- ris, PortJefferson, NY, 1988, unpubl.), we found virtually the same solution as Begle (his pub- lished results are in square brackets): four short- est trees [Begle reported two], length 215 steps [215], with the same strict consensus (Fig. 1) as Begle's (1992, fig. 2) published tree, and with C.I. 0.54 [0.56] and R.I. 0.82 [0.83] when the 11 autapomorphic characters are omitted (25, 27, 32, 41, 42, 46, 48, 49, 84, 87, and 96). The minor differences in tree statistics could be due to different treatment of multistate characters or polymorphism: we ran all but character 65 as unordered; with all characters unordered, we found the same four trees, length 213 steps. In the analyses described above, polymorphisms for states (0) and (1) were treated as (0). With polymorphisms treated as (?) and character 65 ordered, we found the same four trees, length 213, C.I. 0.55 and R.I. 0.82 with autapomorphies omitted; or if all characters are unordered length 211, C.I. 0.56, R.I. 0.82 with autapomor- phies omitted. The virtual coincidence between these results and Begle's shows that our com- plaints do not merely concern misprints in Be- gle's published matrix. When we ran the corrected matrix (see Ap- pendixes), not surprisingly we found a very dif- ferent result. There are three equally parsimo- nious trees with the strict consensus in Figure 2. Comparison of Figures 1 and 2 shows that, of the 14 nodes in Figure 1, only three (less than a quarter) are reproduced in Figure 2 (num- bered 1-3). Major differences between Figures 1 and 2 include the following: (1) the position of esocoids, which are in the basal polychotomy in Figure 1 and are the sister group of osme- roids in Figure 2; (2) Argentinoidei (the node beyond Osmeroidei in Fig. 1) are nonmono- phyletic in Figure 2, with the argentinoid gen- era (Argentina, Glossanodon, Bathylagus, and Op- isthoproctus) placed as the sister group of ale- pocephaloids + esocoids + osmeroids; (3) Ale- pocephaloidea are nonmonophyletic in Figure 2, with Bathylaco placed as the sister group of esocoids + osmeroids + the remaining alepo- cephaloids; and (4) relationships within the re- maining alepocephaloids (Narcetes and the taxa distal to it in Fig. 2) are totally incongruent with the pattern in Figure 1. The differences be- tween Figures 1 and 2 demonstrate the effects of Begle's miscoding, but in our opinion, the corrected version (Fig. 2) does not have much to do with the relationships of the taxa con- cerned (Fig. 3); Figure 2 is more a reflection of Begle's choice of characters. For example, the characters that place esocoids with osmeroids in Figure 2 are absence of the orbitosphenoid, me- socoracoid, and pubic symphysis and reduction of the articular and the dorsal portion of the opercle; in the context of a wider survey of low- er euteleostean characters, these are not con- vincing evidence that the two groups are im- mediately related. ? 1997 by the American Society of Ichthyologists and Herpetologists 402 COPEIA, 1997, NO. 2 utgroups Salmonoids Esocoids Neoteleostei LEI -osmeroidei Glossanodon Argentina Opisthopraoctus Lepochilichthys Bathylaco Narcetes Rouieina lAepocephalus Badhyoono Talismania - Platytroctidae Fig. 1. Begle's (1992, fig. 2) tree of lower eute- leosts, based on his data (Appendixes). Strict consen- sus of four shortest trees (length 215 steps; see text for details) found with Hennig86 (vers. 1.5, J. S. Far- ris, Port Jefferson, NY, 1988, unpubl.). As in Begle's published tree, the 15 taxa of osmeroids (Galaxiidae to Lepidogalaxias in Appendix 1) are collapsed into a single node, Osmeroidei. Comments on our recoding.-We discard 19 of the 108 characters in Begle's matrix. The majority (14) of them are among characters 1-84 from Begle (1991), and our reasons for discarding them are summarized in Patterson andJohnson (1997). Among the new characters introduced in Begle's 1992 paper, we discard six (83, 94, 98, 99, 100, and 105). Number 83 concerns the al- veolar process of the premaxilla and is discard- ed because the supposed primitive state is apo- morphic for a subgroup of salmonoids (Salmon- inae). Number 94 is redundant, concerning fea- tures of the opercle dealt with under numbers 65 and 82. Numbers 98 and 99 concern the lat- eral ridge or strut on the hyomandibular. Num- ber 98 mistakes the primitive state (a long ridge) for a derived state and codes as primitive a condition already coded (number 26) as de- rived. Number 99 is redundant in part (repeat- ing 26) and self-contradictory in coding. Num- ber 100 concerns laminar bone on the anterior margin of the hyomandibular and is discarded because we could not confirm differences be- tween taxa. Number 105 concerns the opercular process of the hyomandibular and is discarded for the same reason as number 100. In Begle's (1991) matrix, there were obvious warnings on the quality of the data in entries for undescribed states of several characters. The warnings in Begle (1992) are less obvious be- cause they demand some knowledge of fishes. Three examples of errors in characters intro- duced in the 1992 paper are numbers 95, 101, and 108. Under character 95, presence or ab- sence of maxillary teeth, a toothed maxilla was incorrectly assigned by Begle to esocoids; to the utgroups eoteleostei Salmonoids Glossanodon j 2-1-Argentina L-3 -Opisthoprctus Rouleina L|jLeptoderma ETalmiania 1 Aiteceprhao Platytroctidae 1 7 Leptochiichthys N1arcets I -Esocoids 7 smeroidei Fig. 2. Tree for the same taxa as Figure 1 after correction of Begle's data, as in Appendix 1. Strict consensus of three shortest trees (length 232 steps; C.I. 0.42, R.I. 0.73) found with Hennig86 (vers. 1.5, J. S. Farris, PortJefferson, NY, 1988, unpubl.). Nodes common to this tree and Figure 1 are numbered. As in Figure 1, the 15 taxa of osmeroids (Galaxiidae to Lepidogalaxias in Appendix 1) are collapsed into a sin- gle node, Osmeroidei. -Outgroup -Esocoids -Neoteleostei -Bathylagus -Opisthoproctus -Argentina -Glossanodon -Platytroctidae -Bathylaco -Bathytroctes -Narcetes -Talismania -Alepocephalus -Leptochilichthys -Bathyprion -Rouleina -Leptoderma -Salmonoids -Prototroctes -Retropinna -Stokellia -Aplochiton -Galaxiidae -Lovettia -Lepidogalaxias -Hypomesus -Plecoglossus -Mallotus -Salangidae -Osmerus -Allosmerus -Spirinchus -Thaleichthys Fig. 3. Relationships of the taxa in Figures 1 and 2 as inferred by Johnson and Patterson (1996). Tree produced with Clados (vers. 1.2, K. C. Nixon, Tru- mansburg, NY, 1992, unpubl.). PATTERSON AND JOHNSON-COMMENTS ON BEGLE 403 retropinnids Prototroctes and Stokellia; and to Aplochiton, Lovettia, Lepidogalaxias, and galaxiids. Under character 101, gillrakers toothed or toothless, the primitive state was mistakenly taken to be toothless gillrakers; and the rakers of outgroups, salmonoids, neoteleosts, Retropin- na, and esocoids were incorrectly coded as toothless. Under character 108, first preural centrum (PU1) and first ural centrum (U1) in- dependent or fused, the derived fusion was cod- ed by Begle only in Bathylagus and Opisthoproctus (in which it does not occur); the fusion univer- sally found in galaxiids, neoteleosts, osmerids, salangids, retropinnids, etc., was miscoded. One could quibble about whether the true distribu- tion of these three characters is general knowl- edge among ichthyologists. Nelson (1984), the standard text at the time of publication of Begle (1992), records the toothless maxilla in eso- coids, Prototroctes, Aplochiton, Lovettia, and galax- iids, as did Boulenger (1904) for esocoids, Aplo- chiton, and galaxiids. That teleostean (or actin- opterygian, or osteichthyan) gillrakers are prim- itively toothed should be obvious to anyone who has examined the gill arches in (say) Amia, Elops, and Salmo. As Nelson (1969:486) put it: "In Recent fishes there is a complete transition between ordinary tooth plates and well-devel- oped gillrakers, and there can be no doubt that gillrakers are little more than modified plates." Fusion between PU1 and U1 is a little more ob- scure, but since the condition was first distin- guished, its distribution in lower euteleosts has been recorded repeatedly (e.g., Gosline, 1960; Weitzman, 1967; Rosen, 1974; Fujita, 1990) and should be familiar to anyone working in the os- teology or higher-level systematics of these fish- es. In our view, these errors should have been caught by someone during the procedures be- tween Begle drafting his manuscript and its publication in the professional journal of ich- thyologists. Other errors in Begle's matrix are more tech- nical. A typical example taken from the char- acters introduced in Begle's 1992 paper is num- ber 87, branchiostegal cartilages connecting the branchiostegal rays with the hyoid arch. These were coded by Begle as present only in the ale- pocephalid Talismania but had previously been illustrated by Sazonov (1986, figs. 6-7) in seven genera of platytroctid alepocephaloids, and we found them also in the alepocephalids Bajacal- ifornia, Bathylaco, Bathytroctes, Leptochilichthys, Rinoctes, and Alepocephalus tenebrosus and in the osmerids Mallotus and Hypomesus olidus. Many similar examples could be given. Character 71 was a particular problem. Begle's original de- scription concerned a ventral process on the pterosphenoid bone. On investigation, we had to delete the character and replace it by four two-state characters (71A-D in Appendix 1). The first concerns presence or absence of the pterosphenoid: it is absent in Lovettia and sal- angids, which must therefore be coded (?) for any feature of the bone. The other three char- acters (71B-D) concern three different, non- homologous processes on the pterosphenoid (one dorsomedial, one ventral, and one pos- teroventral, directed toward the prootic). Statistics.--As shown in the Appendixes, accord- ing to our reading of the specimens and litera- ture, Begle's (1992) matrix contains errors in 92 (85%) of his 108 characters, leaving only 16 that we accept as he coded them. Of those 16, six (32, 46, 48, 49, 84, and 96) are autapomorphic, leaving only 10 that can be used to group taxa. And those 10 are all taken from previous cla- distic analyses (7, 9, 19, 21, and 54 from Fink, 1984; 30 and 51 from Howes and Sanford, 1987a, 1987b; 53 from Rosen, 1974; and 80 and 102 from Greenwood and Rosen, 1971). A more accurate way of assessing Begle's er- ror rate is to compare the number of asterisks in Appendix 1, each an error, with the total number of entries. Excluding 12 deleted char- acters (62-64, 67, 71, 74, 77, 94, 98-100, and 105) in which errors cannot be assessed for one reason or another, the total entries in the ma- trix are 32 X (108 - 12) = 3072. There are 382 asterisks in Appendix 1, giving an error rate of 12.5%. A more telling way of assessing Begle's error rate is to compare the number of asterisks with the number of positive (nonzero) entries in the original matrix, since only the positive entries comprise Begle's character information. Excluding the 12 problematic deleted charac- ters, there are 657 positive entries in Begle's ma- trix; the 382 asterisks in Appendix 1 give an er- ror rate of 58%. Begle's conclusions on euteleostean interrelation- ships.-Begle (1992) summarized his conclu- sions on euteleostean interrelationships in a tree with six characters (Fig. 4). Two characters (numbers 5 and 6 in Fig. 4) link Begle's Os- merae (alepocephaloids, argentinoids, osme- roids) with Neoteleostei; they are his characters 83 and 100. Number 83, "presence of alveolar process of premaxilla," is incorrectly coded and characterizes all taxa in the diagram except for salmonine salmonoids and Opisthoproctus (which has no premaxilla). Number 100, "reduction in laminar bone on anterior margin of hyoman- dibula," is a character that we discarded be- cause we could not confirm differences between 404 COPEIA, 1997, NO. 2 Outgroup Esocae 1 2 3 Ostariophysl -4?Salmonidae 5 6 Osmerae Neoteleostei Fig. 4. Begle's (1992) interpretation of the rela- tionships of lower eutelosts (after Begle, 1992, fig. 9). Tree produced with Clados (vers. 1.2, K. C. Nixon, Trumansburg, NY, 1992, unpubl.). taxa. One character (number 4) links ostario- physans and salmonoids with Osmerae + Neo- teleostei, distinguishing them from esocoids; it is "loss of toothplate on fourth basibranchial" and is credited to Rosen (1974). But Rosen (1962, fig. 13; Rosen and Patterson, 1969, pl. 65, fig. 1) illustrated this toothplate in the percop- siforms Amblyopsis and Percopsis (it also occurs in Aphredoderus; Nelson, 1969, pl. 92, fig. 2), and Rosen (1974:273) reported it as present in the ostariophysan Ichthyborus and in Lepidogalaxias (cf. Fink and Weitzman, 1982:81; we did not find the toothplate in Lepidogalaxias, which is unusual in having the fourth basibranchial fully ossified). Among argentinoids, the toothplate also occurs in the primitive bathylagid Bathylag- ichthys (Kobyliansky, 1986, fig. 11) and the prim- itive microstomatid Nansenia (Kobyliansky, 1990, fig. 8; pers. obs.). Thus the fourth basi- branchial toothplate occurs in several euteleos- tean groups, and its absence cannot have the significance implied in Figure 4. Three charac- ters in Begle's tree (1-3) link esocoids (Esocae of Rosen [1974] and Begle [1992]) with ostar- iophysans, salmonoids, and Osmerae + Neote- leostei; two are Begle's characters 28 (presence of adipose fin) and 33 (presence of nuptial tu- bercles), and the third, credited to Greenwood and Rosen ("1977", 1971 intended), is "mem- branous outgrowth of first uroneural." Esocoids have no adipose fin (e.g., Boulenger, 1904; Nel- son, 1984); Begle mistakenly coded them (and all sampled alepocephaloids) as having one (Appendix 1, number 28). Esocoids do not de- velop nuptial tubercles (Collette, 1977); Begle mistakenly coded them as having them (Appen- dix 1, number 33). Esocoids do have a mem- branous outgrowth on the first uroneural, but argentinoids (Greenwood and Rosen, 1971) and ostariophysans (Johnson and Patterson, 1996) do not. In short, not one of the charac- ters in Begle's diagram (Fig. 4) has the distri- bution indicated. As with Begle's matrix, the warnings are clear (adipose fin and nuptial tu- bercles reported in esocoids) and should have been caught before publication. Conclusions.-Teleostean fishes are "the verte- brate group to which cladistic analysis was first applied, and, for that reason, teleost systematics has probably progressed further down the road from traditional phenetics towards a phyloge- netic system than has the classification of any other vertebrate group" (Patterson, 1993:621). Begle's (1992) paper is a warning of how the application of cladistics in teleosts can lead the way toward chaos. One consequence of the cla- distic revolution and the development of nu- merical cladistics is that a matrix and a com- puter-generated parsimony analysis are now the norm; if you do not provide them, referees will demand them. The larger the matrix (in num- ber of taxa and of characters), the more im- pressive it is, but the greater is the likelihood that error will creep in. The advantage of re- quiring a matrix is that it presents data in un- ambiguous format, so that they may be checked. However, to check a large matrix as thoroughly as we checked Begle's requires resources: ade- quate material, technical knowledge, time, and above all will power or commitment. It may nev- er have been done before, and we hope it never has to be done again. Those preparing similar matrices might take a hint from molecular sys- tematists, who routinely deal with character data in quantity: "Each sequence [read 'character'] was read and entered twice by two different per- sons" (Le et al., 1993:32). In other words, get some help. Those supervising, editing, or refe- reeing works containing large matrices could use the same advice. Their responsibilities are heavy, because observations published in pro- fessional journals are inevitably regarded as hav- ing been prepared and vetted with sufficient care to approximate truth. Acknowledgments.-For comments on a draft, we are grateful to C. Baldwin, J. Coddington, S. Jewett, L. Parenti, V. Springer, and S. Weitzman. LITERATURE CITED AHLSTROM, E. H., H. G. MOSER, AND D. M. COHEN. 1984. Argentinoidei: development and relation- ships. Am. Soc. Ichthyol. Herpetol. Spec. Publ. 1: 155-169. BEGLE, D. P. 1991. Relationships of the osmeroid fish- es and the use of reductive characters in phyloge- netic analysis. Syst. Zool. 40:33-53. - . 1992. Monophyly and relationships of the ar- gentinoid fishes. Copeia 1992:350-366. PATTERSON AND JOHNSON-COMMENTS ON BEGLE 405 BOULENGER, G. A. 1904. Systematic account of Te- leostei, p. 541-727. In: Fishes, ascidians, etc. The Cambridge natural history. Vol. 7. S. F. Harmer and A. E. Shipley (eds.). Macmillan, London. CHAPMAN, W. McL. 1942. The osteology and relation- ships of the Argentinidae, a family of oceanic fishes. J. Wash. Acad. Sci. 32:104-117. COLLETTE, B. B. 1977. Epidermal breeding tubercles and bony contact organs in fishes. Symp. Zool. Soc. Lond. 39:225-268. FINK, W. L. 1984. Basal euteleosts: relationships. Am. Soc. Ichthyol. Herpetol. Spec. Publ. 1:202-206. - 9, AND S. H. WEITZMAN. 1982. Relationships of the stomiiform fishes (Teleostei), with a description of Diplophos. Bull. Mus. Comp. Zool. 150:31-93. FUJITA, K. 1990. The caudal skeleton of teleostean fishes. Tokai Univ. Press, Tokyo, Japan. GOSLINE, W. A. 1960. Contributions toward a classi- fication of modern isospondylous fishes. Bull. Br. Mus. (Nat. Hist.), Zool. 6:325-365. GREENWOOD, P. H., AND D. E. ROSEN. 1971. Notes on the structure and relationships of the alepocephal- oid fishes. Am. Mus. Novit. 2473:1-41. HowEs, G. J., AND C. P. J. SANFORD. 1987a. Oral on- togeny of the ayu, Plecoglossus altivelis and compar- isons with the jaws of other salmoniform fishes. Zool. J. Linn. Soc. 89:133-169. - , AND - . 1987b. The phylogenetic posi- tion of the Plecoglossidae (Teleostei, Salmonifor- mes), with comments on the Osmeridae and Os- meroidei. Proc. 5th Congr. Europ. Ichthyol., Stock- holm 1985:17-30. JOHNSON, G. D., AND C. PATTERSON. 1996. Relation- ships of lower euteleostean fishes, p. 251-332. In: Interrelationships of fishes. M. L. J. Stiassny, L. R. Parenti, and G. D. Johnson (eds.). Academic Press, San Diego, CA. KOBYLIANSKY, S. H. 1986. Materials for a revision of the family Bathylagidae (Teleostei, Salmonifor- mes). Trudy P. P. Shirsov Inst. Oceanol. 121a:6-50 (In Russian). . 1990. Taxonomic status of microstomatid fishes and problems of classification of suborder Ar- gentinoidei (Salmoniformes, Teleostei). Ibid. 125: 148-177 (In Russian). LE, H. L. V., G. LECOINTRE, AND R. PERASSO. 1993. A 28S rRNA-based phylogeny of the gnathostomes: first steps in the analysis of conflict and congruence with morphologically based cladograms. Mol. Phy- log. Evol. 2:31-51. MCALLISTER, D. E. 1968. Evolution of branchiostegals and classification of teleostome fishes. Bull. Nat. Mus. Can. 221:1-239. NELSON, G. J. 1969. Gill arches and the phylogeny of fishes, with notes on the classification of verte- brates. Bull. Am. Mus. Nat. Hist. 141:475-552. NELSON, J. S. 1984. Fishes of the world. 2d ed. John Wiley and Sons, New York. PATTERSON, C. 1993. Teleostei, p. 619-654. In: The fossil record 2. M. J. Benton (ed.). Chapman and Hall, London. ,, AND G. D.JOHNSON. 1997. The data, the ma- trix and the message: comments on Begle's "Rela- tionships of the osmeroid fishes." Syst. Biol. 40. ROSEN, D. E. 1962. Comments on the relationships of the north American cave fishes of the family Am- blyopsidae. Am. Mus. Novit. 2109:1-35. - . 1974. Phylogeny and zoogeography of sal- moniform fishes and relationships of Lepidogalaxias salamandroides. Bull. Am. Mus. Nat. Hist. 153:265- 326. ,- , AND C. PATTERSON. 1969. The structure and relationships of the paracanthopterygian fishes. Ibid. 141:357-474. SAZONOV, Y. I. 1986. Morphology and classification of the fishes of the family Platytroctidae (Salmonifor- mes, Alepocephaloidei). Trudy P. P. Shirsov Inst. Oceanol. 121a:51-96 (In Russian). WEITZMAN, S. H. 1967. The origin of the stomiatoid fishes with comments on the classification of sal- moniform fishes. Copeia 1967:507-540. (CP) Department of Palaeontology, Natural History Museum, London SW7 5BD, England; and (GDJ) Division of Fishes, National Museum of Natural History, Washington, DC 20560. Sub- mitted: 7 May 1996. Accepted: 18 Sept. 1996. Section editor: R. Winterbottom. 406 COPEIA, 1997, NO. 2 APPENDIX 1. BEGLE'S (1992, APPENDIX) MATRIX OF 108 CHARACTERS IN 33 TAXA, AS PRESENTED BY BEGLE AND AS RECODED AFTER CHECKING. We checked characters against specimens where possible or against the literature where specimens are inadequate (e.g., numbers 4, 24) or not at hand (we lacked Bathyprion). There are three rows for each taxon. The upper row is Begle's coding, with (?) for missing data and (B) for polymorphism in states (0) and (1). The second row is our coding, with "X" for deleted characters and bold entries for inferred errors in Begle's coding. The third row summarizes those errors with an asterisk beneath each error and a total at the end. These asterisks do not all coincide with differences between Begle's coding and ours, because in some characters our coding differs through deleting nonexistent states. CHARACTER 7777 111111111 111111111122222222223333333333444444444455555555556666666666771177777777888888888899955555555599 12345678901234567890123456789012345678901234567890123456789012345678901ABCD2345678901234567890123456789012345678 Outgroups 0000000000000000000000000000000000000000000000000000000000000000000000 000000000000000000000000000000000000 Outgroups 000000000000000000X000X0000000000000x000x00XX00x000000000000X0000xxo0000o00000xx 0000000x0x0 xxXox x x0 Salmonoids 000000000000000000000000000000000000000000000000000000?0000000000000000 0000000000000000000000000000000000100 Salmonoids 102000000000000000OXOOOOOOOOO?00000000oo oxoooxooo? 000 00 00xoooooooooxoooxxxoxxx x00 Neoteleosts 00000000000000000000000000000000000000000000000000000110000000000000000 0000000000010000000000000000100000000 Neoteleosts o ooooooooo00000000 0000000000oooxxooooioox000100011 x00000 xx0% xx0 0%0 0 x0xx10 xx000000Xx000x 1 * 3 Galaxiidae 1110111121121112110000?011010000110000110010001000100001111000100111111 0010000000110000000000000000100000100 Galaxiidae 011012112111211201X%000X01B10000100000110B1XOB00010000111100XXX0 X01800XXBOX0000lX0000000000X100XXX1000X101 * * *** * 17 Lovettia 1110111121121112110000?000000000010000110000001000100001111000100000011 0000000000110100000000000000100000100 Lovettia 011012112101211211X000X 0000000001101XX0200010000111100XXX00X000X1??? XX0X0000 lX000001000lXl00XXX1000X101 ,* * * * * 18 Aplochiton 1110111121121112110000?001000000110000110010000000100001111000100111011 0000000000110000000000000000100000100 Aplochiton 211012112111211701X100X00100000010000011001xx0000010000111100XXX01Xl00X01100XX00X00001X0000000000X100XXX10?0X101 , ** 13 Salangidae 1120211101121112000000?000000000110100000010011110001001111000100000010 00?0000000110100000000000000100000400 Salangidae 21212011001121 00X000X 0000000 101007 X121110010001120XXX00X000X1 0XX10X00X010000001X000XXX10X001 ** * * * * * * 27 Retropinna 11101111110000000011111100000100010000000010001000110000110000110000000 0000000000110000000000000020100000100 Retropinna 11100111110100000xiiix1000001000100000011xx100011000011200XXXOOX00X1 XXXOX1OXO XXx000X101 , ** * ** 11 Stokellia 11101111110000000011111100000100010000000010001000110000110000110000001 0000000000110000000000000021100000100 Stokellia 111001111010200000X11X10000010001000000001 100011000011000OioooxxxoOXOOXO1100XX1OxOOo1XO0ooooOOOoX100XXX1000X101 , ** ? * * * * * * * 11 Prototroctes 11101111110000000011111100001000110000000010001000110000110000110000001 0000000000110000000000000020000000100 Prototroctesioooctes 1110ooi1110100000X111X1oioooiiox10xxx00001 10001100001100 OO oxo xiox 1xooixooooooxxx1001 1XX1 0XXX1000X11 , ** , * * * * * * * 11 Plecoglossus 11112000000000000000000001001011011211000001100000000000110001111000001 0100000000010000000000000010000000100 Plecoglossus 011120000000100000X001X00100101101121100000XX0000100000011000XXX10X00X00112XX00X00000X0000001200X000XXX1000X101 Allosmerus 11112000000000000000000001000000011102000001100001000000110111112000001 1010000000011000000000000011100000100 Allosmerus0001120120000000100000X00X001000000011200000XXOOO0xx00000000000100X000X11XXX2X00 IOXO X1XIOOOXOO12XxXX1000 1 ** ** * *9 Hypomesus 11112000000000000000000001000000011202000000100001000000110001111000001 0010000000010000000000000010100000000 Hypomesus B111200000001000000X0010100000001020200000XX0000100000011200XXX 10X00000100XX00X00B00X000000 00X000XXX1000X001 * 10 Osmerus 11112000000000010000000101000010011212000001100000000000110001111000001 0100000000010000000000000010100000100 Osmerus 111120000000100100X000X10100000001120200000XX000010000011200XXX10X000X0001 X00X00000X000000100X000XXX1000X101 , ** ** * 11 Spirinchus 11112000000000000000000001000000011102000001100001000000110111112000001 1010000000010000000000000010100000100 Spirinchus 11112 1 00000001B12002001 0000000112 11 2XXOxX 0000X0OXXX1x0OX101 ** * 9 Mallotus 11112000000000000000000001000000011101000001100001001000110001112000001 0010000000010000000000000010100000000 Mallotus 111120000 000 0 1010200000 0XX000010100011210XXX20X000X00100XX00X00000X00010000X000XXX1000X001 * 10 Thaleichthys 11111000000000000000000100000000011202000001100000000000110100111000001 0000000000010000000000000011100000100 Thaleichthys 111120000000100000X000X10000000001120200000XX0000100000011211XXX10X000X00012XXO0X00000X000000100X000XXX1000X101 , ,** ** 11 Lepidogalaxias 01200111200211001100000000110000100000001110001000102111110000100000111 0001110000110000000000000000100000400 Lepidogalaxias 012003112001210011X000X00111000010000010111XX0200011211101700XXX00X011X00100XX10X00001X0100010000X100XXX1000X400 * * * 16 Esocoids 0000001000000000000000000000000000000000000000000000000000000000000000 0001110000000000000000000000000000000 Esocoids 01200011000 10000X00XO OOOOOOOOOOOOO 1000010000000000XX2002000000OO1 XXX00X000XO000XX11X0000 00000010000XX0000000 ** *** *** * *** ** * 18 Argentina 00201000000100000000000000000000100000000000000001000000000000010000000 0000001111010010110000010000101100310 Argentina 00201000000000000X00X0 00000010000000000XX000100000000000XXX00X00OX0000OXX00X10100X0010110100X100XXX1110X30B ** 8 Alepocephalus 00201001000000020000000000000000100000000000000000000000110000110000000 0000001111110000001101110100110000400 Alepocephalus 002000010000200200X000X00001000010000001 XX0000000000000XXX00X000X00000XX00X12101X00100100lXl00XXX1000X400 * ** ** * 12 Bathylagus 00201010000000000000000000000000100000000000000001000000110000010000000 0000001111010110010110010000101100411 Bathylagus 0020101000002 0000XOOX000000001O0000XX00001000000B0010XXX00X000X00000XX00X1111X0000010000X100XXX1110X401 ** * ** * ** 11 Talismania 0020B001000000000002000000000000100000000000000000000000000000110000000 0000001111110101001101100100110010400 Talismania 002000010000200000X000X00001000010000001000XX0000000000000000XXX00X000X00000XX00X10101X0101001001X000XXX0001X400 Platytroctidae 00208000000000020002000000000000100000000000000000000000BB0000110000000 000000111110100001B01101100110001400 Platytroctidae 001000010000800000 100001000X0X00X00000XX0OX121OIXOB01O00BOOX011XXXO000X10B ** * * * * * * * 18 Leptoderma 0020?011010000120000000000000000100000000000000000000000000000110000000 0000001111110100001101110101110001400 Leptoderma 00200011010101201X000X00001000010000001000XX0000000000000000XXX00X000X00000XX0lX1210lX010000000lX100XX000lX40 1 * * * 14 Leptochilichthys 00201000000000000000000000000000100000000000000000000000100000110000000 0000001111110000000100110100010001?00 Leptochilichthys 00200001000020 000000 0000001000XX0000000 0000011000XXX00X000XO00000XX00X10101X00100100X101XXX0000X?00o * * 11 Opisthoproctus 00201010000000000000000000000000100000000000000001000000110000010000000 0000001111010110110110010000101100401 Opisthoproctus 00200010000 0000X000X0000070001000001010 XX000 X 00XX0X01110X0110010010X100XXX1110X410 * * **14 Bathyprion 00201010000000020000000000000000100000000000000000000000110000110000000 0000000111110000001101100101110010400 Bathyprion 002000110000?00200X000X0000100001000000?000XX0000000000011000XXX00X000X00000XX00X010X000?001001X000XXX0001X400 Bathytroctes 00201000000000020000000000000000100000000000000000000000100000110000000 0000001111110000001100100101110001400 Bathytroctes 002000010000200200X000X00001000010000000000XX0000000000000000XXX00X000X00000XX00X12101X000100100X000XXX0000X400 SC C * * 10 Bathylaco 0020?00000000000000000000000000010000000110000000 0000000111110000000100200101110010400 Bathylaco 002000010000000000XOXOOOO10000100 0 0X 0 0XXOOOOOOOOOOO XXX1OXOOXO OXXXOO11 OX000100000XOOOXXX) 00X4001 Glossanodon 00201000000100000000000000000000100000000000000001000000100000010000000 0000001111010B10110000010000101100210 Glossanodon 002010000000000000X000X00000000010000000000XX0000100000000000XXXOOXOOOX00000XX00X10100XOB10110BOOX100XXX 111X20B * * ** 9 Narcetes 00201000000000000000000000000000100000000000000000000000110000110000000 0000001111110000000100100101010000400 Narcetes O020000100002000X00OOIOOOOOOOOOOOOOOOOOOOXXXOOXOOOXOOOOXX0X1011OIX00?00XOOOXOOctes 200001000020000010000100000010001X )0X400 SRouleina 00201000010000000000000000000000100000000000000000000000110000110000000 0000001111110100000101100101110000400 ;Rouleina 002000010100200200X000X0000100001O000001000XX00000000000 00XXX00X000X00000XX00X10101X0100001001X000XXX0000X400 S C C *** *C* * * * C13 PATTERSON AND JOHNSON-COMMENTS ON BEGLE 407 APPENDIX 2. CHARACTERS, ABRIDGED FROM BEGLE (1992). Comments in square brackets detail changes we made or add explanation; characters 1-82 and 84 were included in Begle (1991) and our brief comments are in Patterson and Johnson (1997). 1. Vomer. 0 = composed of head and posterior shaft; 1 = shaft absent. [State (2) added for absence of vomer in Aplochiton and salangids.] 2. Articular. 0 = fused; 1 = absent or appearing late. 3. Mesopterygoid teeth. 0 = distributed over surface of bone; 1 = narrow band of larger teeth along medial border; 2 = absent. 4. Anchor membrane of egg. 0 = egg not adhesive; 1 = adhesive anchor membrane present. 5. Caudal fusion pattern. 0 = no fusion of rudimentary neural arches to any element in caudal skeleton; 1 = arches fuse first to centrum, followed in some case by fusion to first uroneural; 2 = arches fuse first to uroneural, followed in some case by fusion to centrum. [In Begle (1991) the character was two-state, discriminating osmerids and salangids from other taxa by fusion of first uroneural with centrum.] 6. Orientation of infraorbital sensory canal. 0 = posterodorsally deflected; 1 = posteroventrally deflected. [See also no. 19.] 7. Mesocoracoid. 0 = present; 1 = absent. 8. Dorsal fin position. 0 = anterior; 1 = posterior. 9. Principal caudal rays. 0 = 10/9; 1 = 9/9; 2 = 8/8. 10. Palatine teeth. 0 = present; 1 = absent. 11. Ectopterygoid. 0 = present; 1 = absent. [See also no. 23.] 12. Extrascapular. 0 = present; 1 = attached to pterotic; 2 = absent. [In Begle (1991) the character was two- state; state (1) added in Begle (1992) for Argentina and Glossanodon, with Chapman (1942) as source. Chapman chose to call the dermopterotic of Argentina the supratemporal and described it as fused to the pterotic (= autopterotic), the normal teleostean condition. Begle followed Ahlstrom et al. (1984), not specimens, in interpreting Chapman's supratemporal as the extrascapular. Begle's state (1) is deleted and his (2) replaced by (1).] 13. Cleithrum process. 0 = cleithrum with ventral process descending to meet coracoid; 1 = without ventral process. [Two conditions are conflated under state (0). The primitive state, here coded (0), is a long- based, triangular process against which the coracoid lies. In osmerids and retropinnids, here coded (1), there is a narrow-based, columnar process that interdigitates with the coracoid. Absence is here coded (2).] 14. Posterior pubic symphysis. 0 = present; 1 = absent. 15. Scales. 0 = present; 1 = absent. 16. Vomerine teeth. 0 = present; 1 = "huge fangs"; 2 = absent. [Vomer absent in Aplochiton and salangids, here coded (?).] 17. Posterior border of operculum. 0 = rounded; 1 = incised or emarginate. 18. Number of hypurals. 0 = 6; 1 = 5. 19. Extent of infraorbital canal. 0 = not extending to preopercle; 1 = extending to preopercle. [State (1), in retropinnids, here added to no. 6, as state (2), and this character deleted.] 20. Ceratohyal border. 0 = more or less straight, with branchiostegals along most of its length; 1 = deeply concave anteriorly, with branchiostegals restricted to area posterior to concavity; 2 = ventral border with rectangular notch. [In Begle (1991) the character was two-state. State (2) added in Begle (1992) for Talismania and platytroctids. The notch is in the dorsal border of the ceratohyal, and is homologous with the fenestra ("beryciform foramen" of McAllister, 1968) primitively present in the teleostean ceratohyal.] 21. Horny midventral abdominal keel. 0 = absent; 1 = present. 22. Ovaries. 0 = left and right present; 1 = right only. [It is the left ovary only that is present in retropinnids (and Plecoglossus). ] 23. Location of ectopterygoid. 0 = posterior to autopalatine; 1 = ventral to autopalatine. [The bone is the dermopalatine (no. 11); character deleted.] 24. Cucumber odor. 0 = freshly caught specimens lacking cucumber odor; 1 = fresh specimens with distinc- tive cucumber odor. 25. Basioccipital lateral pegs. 0 = basioccipital without lateral extensions; 1 = with pegs projecting caudally along each side of first vertebra. [Character deleted; see comment in Patterson and Johnson (in press).] 26. Lateral hyomandibular spur. 0 = hyomandibular without lateral projection; 1 = with lateral spur, at or below level of its articulation with opercle. projecting caudally to contact preopercle. 27. Posterior border of caudal fin. 0 = incised or deeply forked; 1 = rounded or emarginate. 28. Adipose fin. O = present; 1 = absent. 29. Articulation of premaxilla with maxilla. O = premaxillary articular process not tightly adhering to max- illary head; 1 = syndesmosis between the two bones. [Premaxilla absent in Opisthoproctus, here coded (?).] 408 COPEIA, 1997, NO. 2 APPENDIX 2. CONTINUED. 30. Palatine-maxilla rticulation. 0 = palatine without articulatory head meeting similar head on maxilla; 1 = maxilla and palatine with head-to-head articulation. 31. Symphysial cartilages. 0 = paired cartilages not present at dentary symphysis; 1 = paired cartilages present at symphysis. 32. Ectopterygoid flange. 0 = dorsal rim of ectopterygoid not directed laterally; 1 = dorsal rim directed laterally in a horizontal flange. 33. Nuptial tubercles. 0 = present; 1 = absent. 34. Pelvic girdle. 0 = without ventral condyle; 1 = with ventral condyle articulating with the first three or four hemitrichs. 35. Autopalatine. 0 = not in the shape of a dumbbell; 1 = in distinctive dumbbell shape. 36. Adipose cartilage. 0 = absent; 1 = present; 2 = present, pear-shaped. 37. Dentary symphysis. 0 = without medial process; 1 = with medial tusk-like process. 38. Metapterygoid. 0 = without lateral shelf; 1 = with short lateral shelf; 2 = with prominent diagonal shelf. 39. Uncinate process on first epibranchial. 0 = present; 1 = absent. 40. Uncinate process on second epibranchial. 0 = present; 1 = absent. 41. Uncinate process on third epibranchial. 0 = present; 1 = absent. 42. Uncinate process on third pharyngobranchial. 0 = present; 1 = absent. 43. Uncinate process on second pharyngobranchial. 0 = present; 1 = absent. 44. Uncinate process of third pharyngobranchial. 0 = not extending over second epibranchial; 1 = extending well over body of second epibranchial. 45. Uncinate process of second pharyngobranchial. 0 = directed laterally or caudally; 1 = directed anteriorly. 46. Hyomandibula. 0 = not fused to palatine [palatoquadrate intended]; 1 = fused with palatine [palato- quadrate]. 47. Third pharyngobranchial. 0 = with narrow anterior extension, reaching first pharyngobranchial; 1 = without anterior extension. 48. Occiput. 0 = not greatly depressed; 1 = greatly depressed. 49. Separate fourth hypobranchial. 0 = absent; 1 = present. 50. Levator process on fourth epibranchial. 0 = wide, its width at distal margin up to half the length of underlying epibranchial; 1 = very narrow, narrower than width of underlying epibranchial. 51. Nasal lamellae. 0 = arranged in rosette; 1 = parallel, longitudinal. 52. Preopercular and supraorbital sensory canal pores. 0 = five or more; 1 = three. 53. Anal rays/scales. 0 = not modified in males; 1 = scales anterior to dorsal margin of anal fin greatly enlarged, anal-fin rays and supports also more or less modified; 2 = as (1) but fin rays and supports extremely modified. [In Begle (1991) the character was two-state; state (2) added in Begle (1992) for Lepidogalaxias alone.] 54. Retractor dorsalis. 0 = absent; 1 = present. 55. Occipital condyle. 0 = formed by basioccipital; 1 = tripartite, formed by basioccipital and exoccipitals. 56. Temporal sensory canal. 0 = present; 1 = absent. 57. Basisphenoid. 0 = present; 1 = reduced or absent. [In Begle (1991) state (1) was absence of the bone. "Reduced" might have been added in Begle (1992) to excuse his (1991) coding of absence in alepoce- phaloids and argentinoids.] 58. Orbitosphenoid. 0 = present; 1 = reduced or absent. [As no. 57 in Begle (1991).] 59. Fusion of fifth epibranchial to fourth. 0 = no; 1 = yes, to form circular foramen for efferent artery. 60. Quadrate. 0 = without linear ridges; 1 = narrow transverse ridges present, sometimes ramifying. 61. Metapterygoid. 0 = without linear ridges; 1 = ridges present, sometimes ramifying. 62. Epihyal. 0 = without foramen; 1 = with midlateral foramen. 63. Pterosphenoid. 0 = not reduced, meeting on midline; 1 = reduced and widely separated, not meeting at midline. 64. First basibranchial. 0 = unmodified; 1 = with ventral cartilaginous vane. 65. Anterodorsal border of opercle. 0 = without spine, horizontal; 1 = with notch and spine; 2 = with deep narrow notch. 66. Ascending process of premaxilla. 0 = knoblike; 1 = sharply triangular. [Premaxilla absent in Opistho- proctus, here coded (?).] 67. Maxilla. 0 = more-or-less traight in lateral profile; 1 = curved dorsally in lateral profile. 68. Epihyal. O = usually greater than half length of ceratohyal; 1 = short, very much less than half length of ceratohyal. 69. Fifth ceratobranchial. O = without bony laminar extensions near medial margin; 1 = with anterior laminar extension close to medial margin. 70. Palatine. O = contacting maxilla by small knob, if at all; 1 = lateral knob overlying maxilla. PATTERSON AND JOHNSON-COMMENTS ON BEGLE 409 APPENDIX 2. CONTINUED. 71. Pterosphenoid. 0 = without ventral processes; 1 = with small ventral flange midway along its length. 72. Anterior margin of prootic. 0 = rounded and smooth; 1 = notched with a small dorsal projection. 73. Prootic/pterosphenoid contact. 0 = at dorsal margin of prootic; 1 = more medial, contact by interdigi- tation. 74. Lateral spine on sphenotic. 0 = blunt; 1 = rodlike. 75. Pyloric caeca. 0 = present; 1 = absent. 76. Uroneurals. 0 = more than one; 1 = one. 77. First preural centrum (PU1). 0 = more than one neural arch over PUl; 1 = a single rudimentary neural arch over PU1. 78. Accessory cartilage at tip of fifth ceratobranchial. 0 = absent; 1 = present. 79. Fourth ceratobranchial. 0 = unmodified; 1 = in dorsal view, much wider than first three ceratobranchials, sometimes with distal end expanded. 80. Fourth gill arch. 0 = unmodified; 1 = with fleshy membrane along joint of ceratobranchial and epibran- chial partitioning crumenal pouch from orobranchial chamber. 81. Gillrakers on fourth and fifth arches. 0 = unmodified; 1 = expanded/elongate. 82. Opercle. 0 = extending dorsally above articulation with hyomandibular; 1 = not extending above artic- ulation. 83. Premaxilla. 0 = without postmaxillary process (premaxilla and maxilla serially arranged); 1 = with post- maxillary (alveolar) process extending beneath maxilla. [In Begle (1991) the character referred to the length of the alveolar process and was coded as autapomorphic for Stokellia. In Begle (1992) the coding distinguishes neoteleosts, argentinoids, alepocephaloids and osmeroids (all with 1) from esocoids, salm- onoids and outgroups. See comments in text.] 84. Midlateral band of silver pigment. 0 = absent; 1 = present. 85. Pelvic splint. 0 = present; 1 = absent. 86. Shape of interhyal. 0 = elongate, rodlike; 1 = short, dumbbell-shaped. 87. Branchiostegal cartilages. 0 = absent; 1 = small cartilages present, connecting branchiostegals with hyoid arch. [See comments in text.] 88. Ventral arm of symplectic. 0 = short, less than half length of dorsal arm; 1 = elongate, longer than dorsal arm. 89. Metapterygoid. 0 = large, broad; 1 = reduced and rodlike. 90. Medial shelf on metapterygoid. 0 = absent; 1 = present. 91. Uncinate process on fourth epibranchial. 0 = present; 1 = absent. 92. Fifth epibranchial. 0 = short, much less than half length of fourth; 1 = long, almost equal in length to fourth. 93. Ventral coracoid process [= postcoracoid process]. 0 = short, not extending below ventral margin of pectoral girdle; 1 = narrowly elongate, extending below pectoral girdle. 94. Dilatator spine on dorsal margin of opercle. 0 = absent; 1 = present; 2 = large, extending dorsally. [Character discarded, see comment in text.] 95. Maxillary teeth. 0 = present; 1 = absent. [See comment in text.] 96. Shoulder organ. 0 = absent; 1 = present. 97. Elevated median basibranchial ridge. 0 = absent; 1 = present, separating right and left portions of branchial basket. 98. Lateral ridge on hyomandibular. 0 = short, less than half length of hyomandibular; 1 = longer, sometimes occupying entire shaft of bone; 2 = absent. [Character discarded, see comment in text.] 99. Lateral ridge on hyomandibular. 0 = contacts preopercle; 1 = does not contact preopercle. [Character discarded, see comment in text.] 100. Laminar extension on anterior margin of hyomandibular. 0 = present; 1 = greatly reduced or absent. [Character discarded, see comment in text.] 101. Teeth on gillrakers. 0 = absent; 1 = present. [See comments in text.] 102. Mouth. 0 = large; 1 = very small. 103. Vomer. 0 = does not extend anteriorly beyond ethmoid cartilage; 1 = extends anterior to ethmoid cartilage. [Aplochiton and salangids, in which the vomer is absent, cannot be assessed for this character and should be coded (?).] 104. Ovarian tunic. O = absent; 1 = ovaries covered more-or-less completely with membranous tunic. 105. Opercular process of hyomandibular. O = straight, dorsally located; 1 = ventrally curved, dorsally located; 2 = located at or below midpoint of hyomandibular. [Character discarded, see comment in text.] 106. Basihyal teeth. O = scattered over basihyal; 1 = fangs on margin; 2 = small teeth on terminus only; 3 = fangs on terminus; 4 = absent. 107. Development of pectoral fins. 0 = small, relatively late in ontogeny; 1 = large, relatively early in ontogeny. 108. Fusion of PU1 to U1. O = no; 1 = yes. [See comments in text.]