Phylogeny of the Genera and Families

of Zeiform Fishes, with

Comments on Their Relationships with

Tetraodontiforms and Caproids

JAMES C.TYLER,

BRUCE O'TOOLE,

and

RICHARD W1NTERBOTTOM

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 618
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SMITHSONIA N CONTRIBUTION S T O ZOOLOG Y • NUMBE R 61 8

Phylogeny of the Genera and Families

of Zeiform Fishes, with

Comments on Their Relationships with

Tetraodontiforms and Caproids

James C. Tyler, Bruce O 'Toole,

and Richard Winterbottom

Smithsonian Institution Press

Washington, D.C.

2003
ABSTRAC T

Tyler, James C, Bruce OToole, and Richard Winterbottom. Phylogeny of the Genera and

Families of Zeiform Fishes, with Comments on Their Relationships with Tetraodontiforms and

Caproids. Smithsonian Contributions to Zoology, number 618, 110 pages, 90 figures, 2

tables, 2003.—This cladistic study, using 103 putatively informative characters (mostly osteological) of representatives of all of the genera of zeiforms, a primitive tetraodontiform, both

genera of caproids, and numerous outgroups, strongly supports the monophyly of zeiforms.

The possible relationship of zeiforms with tetraodontiforms and caproids is ambiguous; it is

supported in three of the four analyses (data ordered with and without most meristic features

and data unordered with most meristic features) but not in the one that we consider the most

rational and best justified (data unordered without most meristic features).

Within zeiforms, there is strong support for a phylogeny that requires a substantially different arrangement of genera within a new concept of six monophyletic families (three with subfamilies) in two suborders.

The genus Cyttus (Cyttoidei, Cyttidae) is shown to be the sister group to all other zeiforms

(Zeioidei). The study indicates that the zeiform families, as recognized herein, have the following phylogenetic sequence: Cyttidae {Cyttus)—Oreosomatidae (Pseudocyttinae, Pseudocyttus;

Oreosomatinae, Allocyttus, Oreosoma, Neocyttus)—Parazenidae (Parazeninae, Parazen; Cyttopsinae, Cyttopsis, Stethopristes)—Zeniontidae {Zenion, Capromimus, Cyttomimus)—

Grammicolepididae (Macrurocyttinae, Macrurocyttus; Grammicolepidinae, Xenolepidichthys,

Grammicolepis)—Zeidae {Zeus, Zenopsis).

The newly composed families and subfamilies are systematically defined, and keys to them

are provided.

Examples of nearly all of the features used in the phylogenetic analyses and systematic

accounts are illustrated.

SERIES COVER DESIGN: The coral Montastrea cavernosa (Linnaeus). OFFICIAL PUBLICATION

DATE is handstamped in a limited number of initial copies and is recorded in the Institution's

annual report, Annals of the Smithsonian Institution.

Library of Congress Cataloging-in-Publication Data

Tyler, James C, 1935-

Phylogeny of the genera and families of Zeiform fishes, with comments on their

relationships with tetraodontiforms and caproids / James C. Tyler, Bruce O'Toole, and

Richard Winterbottom.

p. cm.—(Smithsonian contributions to zoology ; no. 618)

Includes bibliographic references.

1. Zeiformes—Phylogeny. I. O'Toole, Bruce. II. Winterbottom, Richard, 1944- III.

Title. IV. Series.

QLl.S54no. 618

[QL637.9.Z4]

590 s—dc21

[597\64] 2002030869

© The paper used in this publication meets the minimum requirements of the American

National Standard for Permanence of Paper for Printed Library Materials Z39.48—1984.
Contents

Page

Introduction 1

Acknowledgments 3

Methods 4

Terminology 4

Zeiform Procurrent Caudal-Fin Rays 5

Zeid Pelvic Fin 6

Coding Conventions 6

Analytical Protocols 6

Tree Characteristics 7

Meristic Character-State Evolution 12

Illustrations 14

Material 15

Results 16

Phylogeny 16

Characters and Character States 16

Phylogenetic Results 21

Monophyly of Zeiformes 22

Monophyly of Cyttus: Cyttoidei, Cyttidae 23

Monophyly of Cyttus australis and Cyttus traversi 24

Monophyly of Pseudocyttus, Oreosoma, Neocyttus, Allocyttus, Parazen, Cyttopsis, Stethopristes, Zenion, Capromimus, Cyttomimus, Macrurocyttus,

Xenolepidichthys, Grammicolepis, Zeus, and Zenopsis: Zeioidei 24

Monophyly of Pseudocyttus, Oreosoma, Neocyttus, and Allocyttus: Oreosomatidae 25

Monophyly of Oreosoma, Neocyttus, and Allocyttus: Oreosomatinae 25

Monophyly of Allocyttus verrucosus and Allocyttus niger 25

Monophyly of Parazen, Cyttopsis, Stethopristes, Zenion, Capromimus, Cyttomimus, Macrurocyttus, Xenolepidichthys, Grammicolepis, Zeus, and

Zenopsis 25

Monophyly of Parazen, Cyttopsis, and Stethopristes: Parazenidae 26

Monophyly of Cyttopsis and Stethopristes: Cyttopsinae 26

Monophyly of Zenion, Capromimus, Cyttomimus, Macrurocyttus, Xenolepidichthys, Grammicolepis, Zeus, and Zenopsis 26

Monophyly of Zenion, Capromimus, and Cyttomimus: Zeniontidae 27

Monophyly of Capromimus and Cyttomimus 27

Monophyly of Macrurocyttus, Xenolepidichthys, Grammicolepis, Zeus, and

Zenopsis 27

Monophyly of Macrurocyttus, Xenolepidichthys, and Grammicolepis: Grammicolepididae 28

Monophyly of Xenolepidichthys and Grammicolepis: Grammicolepidinae .. . 28

Monophyly of Zeus and Zenopsis: Zeidae 29

Monophyly of Zenopsis conchifer and Zenopsis nebulosus 29

Previous Phylogenetic Analyses of Zeiforms 29

Phylogenetic Summary 34

Systematics 35

Revised Familial Classification of Zeiformes 35

Characterizations of the Families and Subfamilies of Zeiformes 36
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Key to the Families and Subfamilies of Zeiformes 40

Conclusion 46

Appendix: Character-State Transformations for the Unordered Reduced Data Set

Consensus Tree 47

Literature Cited 48

Figures 51
Phylogeny of the Genera and Families

of Zeiform Fishes, with

Comments on Their Relationships with

Tetraodontiforms and Caproids

James C. Tyler, Bruce O 'Toole,

and Richard Winterbottom

Introduction

Zeiformes (dories), such as Zeusfaber (the John Dory or St.

Peter's fish), have been known since ancient times. Linnaeus

(1758) placed Zeus (sensu lato) in his Thoracici, along with

many groups considered today to be acanthopterygians. In this

he was followed by Cuvier (1817) and by Cuvier and Valenciennes (1835), although the latter placed Zeus and Capros in a

chapter separate from those dealing with other acanthopterygians, such as scombroids, lamprids, carangids, and others.

Giinther (1860) erected the Cyttina (Zeus, Cyttus, Oreosoma)

as one of five groups in his family Scombridae (the other four

being the Scombrina, Nomeina, Stromateina, and Coryphaenina). Gill (1863) erected the Zenoidae (equivalent to

Gunther's Cyttina) as distinct from his Scombroidae, and he

proposed two subfamilies, the Zeinae (Zeus, Zenopsis, Cyttus,

Cyttopsis) and the Oreosomatinae (Oreosoma). Gill did not

speculate on the relationship of the Zenoidae and Scombroidae to other families. Starks (1898) concluded that zeids were

related to chaetodontoids and acanthuroids and not to scombroids. Jordan and Evermann (1898) considered the relationship of zeids to other families to be uncertain, with similarities

to both berycoids (based on their increased number of pelvicfin rays) and acanthuroids (based on their close attachment of

the posttemporal to the skull).

James C. Tyler, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560-0106. Bruce O'Toole and Richard

Winterbottom, Centre for Biodiversity and Conservation Biology,

Royal Ontario Museum, 100 Queens Park, Toronto, Ontario. Canada

M5S 2C6, and Department of Zoology, University of Toronto, Toronto,

Ontario, Canada M5S 1A1.

Holt (1894) suggested that zeiforms might be related to the

Heterosomata (flatfishes) (although he also hinted that flatfishes might have other relatives). Thilo (1901) also believed

that the Heterosomata (pleuronectiforms) were related to zeids.

Boulenger (1902), quoting Holt (1894), independently arrived

at the same conclusion as Thilo; thus, he placed the Zeidae

(along with the fAmphistiidae and the Pleuronectidae) in Division III (Zeorhombi) of his Acanthopterygia. Boulenger's definition of the zeids included a few specialized characters, but it

was based mostly on primitive features. He suggested that the

Zeorhombi were derived from a common ancestor with the

Berycidae. In 1904, Boulenger reiterated this view.

Regan (1910) proposed a close relationship between zeiforms (his Zeidae) and caproids (his Caproidae), recognized

them as the order Zeomorphi, and related them to berycoids,

stating that they differed only in certain specializations.

Regan's views were followed by Gregory (1933).

Norman (1934) disagreed with Thilo's (1901) and Boulenger's (1902, 1904) assessment that zeiforms were related to

flatfishes, but he did not present an alternative hypothesis. Ford

(1938) agreed with Norman, noting that the differences in their

vertebral morphology did not support an association between

flatfishes and zeids. Ford commented on the similarity of the

bifurcate first neural arch and spine supporting the first pterygiophore in both Zeus and Capros.

Berg (1940) simply listed his Zeiformes after the berycoids.

He included three families, the Zeidae, Grammicolepididae,

and Caproidae, and stated that caproids bore some resemblance

to chaetodontoids and ephippidids. Myers reviewed the grammicolepidids (1937) and oreosomatids (1960) and made many

preceptive surmises about generic groupings of other zeiforms.

Patterson (1968) regarded zeiforms (including caproids) to

be the apomorphic sister group to berycoids (including stepha-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

noberyciforms), with the two together being the sister group to

percomorphs. Winterbottom (1974) suggested that at least

some zeiforms might be related to tetraodontiforms. Rosen

(1984) provided evidence for a group comprising caproids, tetraodontiforms, and zeiforms, with the latter two being sister

groups (see "Previous Phylogenetic Analyses of Zeiforms," below, for details).

Most recently, Johnson and Patterson (1993) tentatively suggested that zeiforms (excluding caproids) are the sister group to

beryciforms (excluding stephanoberyciforms) plus percomorphs. They went on to state that Zenion and Parazen might be

the more primitive zeiform taxa (morphologically) and provided the only cladistically based statement of relationships

among zeiform families to date. They also offered considerable

evidence for the monophyly of the Zeiformes (see "Previous

Phylogenetic Analyses of Zeiforms" for details); in the present

work we add substantially to that evidence.

Most authors (e.g., Heemstra, 1980, 1986; Tighe and Keene,

1984; Karrer and Heemstra, 1986) presently recognize five

families of zeiforms (or six families if caproids are included

among zeiforms, as in Eschmeyer, 1990, and Nelson, 1994),

usually with the following included genera:

PARAZEN I DAE {Parazen only)

ZENIONTIDAE (either Zenion only or with the enigmatic

Macrurocyttus also included, in which case the family

name Macrurocyttidae is sometimes used)

GRAMMICOLEPIDIDAE {Grammicolepis, Xenolepidichthys)

OREOSOMATIDAE (Allocyttus, Neocyttus, Oreosoma,

Pseudocyttus)

ZEIDAE {Capromimus, Cyttomimus, Cyttopsis, Cyttus, Stethopristes, Zenopsis, Zeus)

Among the genera usually assigned to the Zeidae, the placement of Capromimus and Cyttomimus has been considered

questionable (Heemstra, 1980), although they are accepted in

that family by Bray (1983). In fact, the family Zeidae has become the convenient repository for all of the zeiform genera

that could not be easily placed in any of the other four families.

The genus Daramattus, with two South African nominal species, has sometimes been recognized as distinct from Grammicolepis within the Grammicolepididae (e.g., Heemstra, 1980),

but we agree with Karrer and Heemstra (1986) that the two

species of Daramattus are both synonyms of Grammicolepis

brachiusculus.

We have examined the osteology of representatives of all of

the above valid genera of zeiforms, and most of the species of

zeiforms. The only zeiform species that we at least tentatively

consider to be valid that we have not osteologically examined

are Allocyttus guineensis, valid according to Karrer (1986);

Cyttopsis cypho, valid according to Paxton and Hanley (1989)

and Bray (1983); Zenopsis sp., a new species from the South

China Sea and western Australia being described by Yamada,

Nakabo, and Bray (MS); and several species of Neocyttus, a

genus recently revised by Yearsley and Last (1998), who show

that in addition to the common N. rhomboidalis examined

herein, there are four other valid species, N. acanthorhynchus

(also considered valid by Heemstra, 1980, and Karrer, 1986),

N. helgae, N. psilorhynchus, and an as yet unnamed taxon.

For several other zeiform species we have examined only radiographs or very limited dry skeletal materials, and our data

on these are insufficient for them to be included in our study:

Allocyttus folletti, Zenion leptolepis (either this species has several synonyms, such as longipinnis and japonicus, in the Indowestern Pacific, or there are more than two valid species of

Zenion), Zenopsis oblongus, and Zeus capensis (see "Material").

The nomenclature of the species of zeiforms studied herein is

relatively stable, and we do not list their often lengthy synonymies. For such synonymies, see the publications of P.C. Heemstra, C. Karrer, and D.J. Bray cited above, and the following:

Heemstra (1999), Karrer (1990), Karrer and Post (1990), Quero

(1973), and Wheeler (1973).

The fossil zeiforms are under study by one of us (JCT) and

are not treated herein, mainly because most of the features in

our matrix of phylogenetically informative characters cannot

be determined in the fossils. The three earliest known zeiforms,

however, belong to morphologically primitive genera (see Figure 89 for the Upper Cretaceous Cretazeus) that cannot be

placed in any of the extant families. These are a single species,

Cretazeus rinaldi (Tyler et al., 2001), from the Upper Cretaceous (upper Campanian-lower Maastrichtian) of Nardo, Italy,

about 72 million years ago (m.y.a.), and two species, Protozeus

kuehnei and Archaeozeus skamolensis (both Bonde and Tyler,

in Tyler et al., 2001) from the upper Paleocene-lower Eocene

boundary (Mo-clay, Fur Formation) of Denmark, about 58-59

m.y.a. We have examined the holotype (Instituto Superior Tecnico (1ST), Laboratorio de Geologia Aplicada, Lisboa, 1ST

559, 9 mm SL) of the putative zeiform Palaeocyttus princeps

Gaudant (1977, 1978) from the Middle Cretaceous of Portugal

and agree with Patterson (1993a, 1993b), who noted that it differs substantially from zeiforms, and with Bonde and Tyler

(MS) that it is not a zeiform but is probably a beryciform.

All other zeiform fossils are of Oligocene (about 36 m.y.a.

for the oldest) or younger age and are zeids that can probably

be placed in either the extant Zeus or Zenopsis (see Figure 90

for one of the typical Oligocene species of Zenopsis), and certainly within the Zeus + Zenopsis clade (Baciu, Bannikov, and

Tyler, in prep.).

By comparison, the earliest known tetraodontiforms are a

clade of three Upper Cretaceous taxa (the oldest, from Hakel,

Lebanon, is from about 95 m.y.a.); this clade is the sister group

to the clade comprising all other tetraodontiforms (Eocene to

present, the oldest from about 58 m.y.a.) (Tyler and Sorbini,

1996). Caproids are known only as early as the lower middle

Eocene (Monte Bolca, Italy, about 50 m.y.a.) (Sorbini, 1983;

Sorbini and Bottura, 1988), with taxa that are similar enough to

the extant species that they presently are contained in extant

genera (but at least the earliest caproid, the Monte Bolca Antigonia veronensis Sorbini, lacks a few derived features of the
NUMBER 618

extant species of Antigonia and could be placed in its own genus). We have examined the holotype (Museum National

d'Histoire Naturelle, Paris, HDJ-73-17,20 mm SL) of the putative caproid Microcapros libanicus Gayet (1980a, 1980b) from

the Middle Cretaceous of Lebanon and agree with Zehren

(1989, MS), Bannikov (1991), and Patterson (1993a, 1993b)

that it is a trachichthyid, not a caproid.

All of the taxa of the earliest known caproids, zeiforms, and

tetraodontiforms are relatively much smaller in size than the

fossils of each of these groups from subsequent periods and of

the extant species of each. These earliest taxa, however, are

fully ossified, and the specimens of each apparently are at least

young adults: 21.0 mm standard length (SL) for the Eocene caproid Antigonia veronensis; 15.4-53.1 mm SL for the Upper

Cretaceous zeiform Cretazeus rinaldi; 8.5-10.5 mm SL for the

two upper Paleocene species of zeiforms; and -10-24.5 mm SL

for the three Upper Cretaceous species of tetraodontiforms. This

leads us to suspect that paedomorphic processes were important

in the early evolution of all three groups (Tyler and Sorbini,

1996, had suggested this previously for tetraodontiforms alone).

The primary purpose of the present study is to elucidate the

intrarelationships of zeiform fishes, and the characters selected

for inclusion in the analyses reflect this (see "Analytical

Protocols"). In order to evaluate the relationships between zeiform taxa, we needed to postulate outgroups of the zeiforms for

purposes of character evaluation. We have examined representatives of most of the higher-level groups that have been suggested as possible sister groups of zeiforms. We did not include

pleuronectiforms, a group that seems to us an unlikely candidate, but this possibility will be the topic of a subsequent study.

We evaluate the evidence supporting zeiform monophyly, revisit the question of caproid, zeiform, and tetraodontiform relationships, and propose a hypothesis of the intrarelationships of

zeiform taxa based on a cladistic evaluation of the osteological

features of 20 zeiform and 10 outgroup species. We then compare our hypotheses with the results of various other previous

studies. Finally, we systematically define all of the familial and

subfamilial categories of zeiforms recognized herein.

This study had its origins in 1992, when two of us (JCT and

RW) and G. David Johnson of the Smithsonian Institution

agreed to attempt a brief preliminary analysis of zeiform relationships for possible inclusion in a forthcoming volume on

fish phylogeny. Because of the press of time, that preliminary

work used a single exemplar for each of the then-recognized

five families of zeiforms, and the results were deemed insufficient for publication; however, some of the phylogenetically

interesting features of zeiforms described herein were first discovered in conjunction with G David Johnson.

ACKNOWLEDGMENTS

For loans and gifts of material, or facilitation of examination

of specimens at their institutions, we would like to thank the

following: B.A. Brown, G.J. Nelson, and C.L. Smith, American

Museum of Natural History; J.R. Paxton, D.J. Bray, and M.

McGrouther, Australian Museum; W.N. Eschmeyer and T. Iwamoto, California Academy of Sciences; L.F. Palmer, S.J.

Raredon, S. Smith, and S.L. Jewett, Smithsonian Institution;

E.B. Bohlke, W.G. Saul, J.G. Lundberg, and M. Littmann,

Academy of Natural Sciences of Philadelphia; B. Chernoff and

M.A. Rogers, Field Museum of Natural History; N.V. Parin,

P.P. Shirshov Institute of Oceanology, Russian Academy of

Sciences; K.E. Hartel, Museum of Comparative Zoology, Harvard University; G.K. Yearsley, CSIRO Marine Research, Tasmania; M.E. Anderson and P.C. Heemstra, J.L.B. Smith Institute, Rhodes University; A.F. Bannikov and N.N. Parin,

Paleontological Institute, Russian Academy of Sciences; K.

Amaoka and CO. Nyako, Hokkaido University; T. Nakabo,

Kyoto University Museum; I. Nakamura, Fisheries Research

Station, Kyoto University; L. Sorbini, A. Aspes, R. Zorzin, and

A. Vaccari, Museo Civico di Storia Naturale di Verona; C. Sorbini, Universita di Pisa; C. Karrer, Universitat Hamburg; H.

Furrer, Palaontologisches Institut und Museum der Universitat

Zurich; D.-S. Baciu, Muzeul de Stiinte ale Naturii, Piatra

Neamt; B. Seret, D. Goujet, and J.-P. Barrieu, Museum National d'Histoire Naturelle; N. Bonde, Kobenhavns Universitet;

Colin Patterson, The Natural History Museum, London; E.

Swidnicka, Wroclaw University; T. Kolar-Jurkovsek, Geoloski

zavod Ljubljana; A. Mouraz Miranda, Instituto Superior Tecnico, Lisboa; D. Nolf, Institut Royal des Sciences Naturelle de

Belgique; R.H. Rosenblatt, Scripps Institute of Oceanography;

and F. Santini, Royal Ontario Museum.

Most of the radiographs used herein were taken by S.J.

Raredon, and photographs of entire cleared and stained specimens from which lateral view illustrations were prepared were

taken by V. Krantz and C.C. Hansen, all of the Smithsonian Institution.

D.M. Tyler, Smithsonian Institution, provided much editorial

advice and corrections to various drafts of the manuscript, and

she prepared all of the illustrations for proper size and labeling.

W.A. Recher, Montgomery Village, Maryland, electronically

scanned and produced the final, labeled versions of the illustrations.

W.N. Eschmeyer, California Academy of Sciences, greatly

aided us in our search for the earliest use of family-level names

within zeiforms.

F. Santini, Royal Ontario Museum, reanalyzed the morphological data set in Wiley et al. (2000) and drew our attention to

the confusion caused by the mislabeling of figure 8 in that paper; he also offered many useful comments on our cladistic

analyses.

We have made extensive use of the dry skeletal materials at

the American Museum of Natural History, and we gratefully

acknowledge the efforts of B.A. Brown and GJ. Nelson to curate and expand that extraordinarily useful collection.

During the course of this study, we received much helpful

advice on a wide variety of relevant research matters from col-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

leagues at the Smithsonian Institution: C.C. Baldwin, B.B. Collette, G.D. Johnson, L.R. Parenti, R.P. Vari, and S.H. Weitzman.

We are especially indebted to GD. Johnson for the many insights he has offered us about zeiform relationships, which are

based on his own research and that in conjunction with the late

C. Patterson, as well as on an unpublished preliminary analysis

of zeiform phylogeny in which he and two of us (JCT and RW)

were involved several years ago. He has continued to cooperate

with us during the course of the greatly expanded work leading

to the present publication, and he has also made his extensive

collection of cleared and stained zeiforms and related groups

freely available to us. We thank K. Doyle, Royal Ontario Museum/University of Toronto, for the help he gave us in that

same preliminary analysis, which was based on only five zeiform species, and for his help in this more comprehensive

study. We were encouraged to greatly expand our coverage of

zeiform taxa and to add many more outgroups during the review of the preliminary analysis by J.A. Moore, Yale University, and V.G. Springer, Smithsonian Institution. We followed

their advice, and now, some years later, present our far more

complete work.

The present manuscript benefited greatly from the comments

offered during the formal peer review process by GD. Johnson

and S.H. Weitzman, Smithsonian Institution; M.F. Mickevich,

National Marine Fisheries Service; and an anonymous reviewer.

This study was supported in part by a National Science and

Engineering Research Council Research Grant (OGP0007619)

to RW. This is Contribution Number 12 of The Centre for

Biodiversity and Conservation Biology of the Royal Ontario

Museum.

Methods

TERMINOLOGY.—All lengths are standard length (SL) unless

otherwise stated, and these are given to the nearest whole millimeter (mm), except for the larval stages discussed in the following section, which are given in tenths of a millimeter. CI=

consistency index and RI=retention index (capitalized when

referring to overall tree statistics; lower-case when referring to

individual character statistics). HPU=haemal spine of preural

centrum, PB=pharyngobranchial, PU=preural centrum, NPU=

neural spine of preural centrum, UP=upper pharyngeal toothplate.

We use Arabic numbers for the characters and character

states in our analyses, and (in brackets) Roman numbers for

characters discussed from the literature (see "Previous Phylogenetic Analyses of Zeiforms").

We use the terms Zeiformes and zeiforms for the five families most commonly assigned to that order, herein expanded to

six families, with zeiforms not including caproids (or tetraodonti forms).

The term Zeomorphi is ambiguous, and we do not use it

herein. As proposed by Regan (1910), the order Zeomorphi included two families, the Zeidae for all zeiforms and the Caproidae for Capros and Antigonia. Myers (1937, 1960) followed

Regan in considering the Zeomorphi to include both zeiforms

(with five families) and caproids. Greenwood et al. (1966) recognized the Zeiformes (with Zeomorphi as a synonym) with

the traditional five families of zeiforms plus caproids, and the

inclusion of caproids with zeiforms was followed by Eschmeyer (1990) and Nelson (1994). Rosen (1984), however, used

Zeomorphi as a division comprising the five families of zeiforms, which he proposed as the sister group to the division

Plectognathi (i.e., the traditional tetraodontiform fishes), with

the Zeomorphi (=zeiforms) + Plectognathi (=tetraodontiforms)

clade as the sister group to the family Caproidae, and with the

caproids + Zeomorphi + Plectognathi constituting the order Tetraodontiformes, a very different usage than traditional for the

terms tetraodontiform and zeomorph. Zehren (1987) followed

Rosen (1984) in the use of Zeomorphi = zeiform, but others

have not. Stiassny and Moore (1992) used "zeoid" for the zeiform lineage within a higher category of a problematic "zeiform radiation" that included caproids, zeoids = zeiforms sensu

stricto, and plectognaths = tetraodontiforms. The term Zeomorphi remains available for some higher category that includes

the Zeiformes.

If a species is of a monotypic genus or if a genus has only a

single species examined in this work, that taxon is often referred to by the generic name alone.

Length of the upper jaw is measured from the midline border

of the premaxilla to the ventral end of the maxilla.

The distinction between the abdominal and caudal series of

vertebrae is relatively clear in most zeiforms, with the first caudal vertebra having a much longer haemal spine than those preceding it, and it supports along its anterior border the first one

or more anal-fin pterygiophores. The distinction, however, is

more subjective in some zeiforms, especially in zeids and oreosomatids, because the haemal spines of several vertebrae overlap the lateral surface of the proximal end of the large first

anal-fin pterygiophore, and these haemal spines only gradually

increase in length. In these cases, the first caudal vertebra is

considered to be that in which the haemal spine is alongside the

rear half of the lateral surface of the upper end of the first analfin pterygiophore, and which is followed by the much longer

haemal spine of the second caudal vertebra, with numerous

anal-fin pterygiophores between the short haemal spine of the

first caudal vertebra and the longer spine of the second (see lateral-view illustrations of the entire skeletons of an oreosomatid

and a zeid, Figures 16, 73). For tetraodontiform fishes we retain herein the traditional distinction between abdominal and

caudal vertebral series, which is based on the first caudal vertebra being the first in the series with a much longer haemal

spine than those preceding it, even though it apomorphically

supports the first anal-fin pterygiophore along its posterior border rather than its anterior border as in zeiforms, caproids, and

most other fishes.
NUMBER 618

The caudal peduncular vertebrae are defined as all those, including the urostylar centrum, posterior to the last vertebra

with neural or haemal spines supporting dorsal- or anal-fin

pterygiophores.

For the specialized scales found in some zeiforms, we follow

the currently used terminology of "buckler" for the greatly enlarged (many times larger than the other scales on the body)

and thickened scale plates found variously along the dorsaland anal-fin bases and along the ventral midline from the isthmus to the anus in genera of several families, and "scutes" for

the only slightly enlarged (no more than about twice the size of

the surrounding scales) modified scales with a zipper-like overlapping arrangement along the ventral midline in the Cyttidae.

We have no evidence about any possible developmental or phylogenetic differences between the scales in these convenient

morphological categories.

The terms interneural space and interhaemal space are used

in the revised sense of Birdsong et al. (1988), Baldwin and

Johnson (1993), and Bannikov and Tyler (1995), with the first

space being that between the first and second neural or haemal

spines, and the preneural and prehaemal spaces being those in

front of the first neural and haemal spines.

Statements about ribs (pleural ribs) and epineural bones are

based on ossified elements and do not include what are sometimes ligamentous elements (for which see Patterson and

Johnson, 1995). We note that all zeiforms lack ossified ribs on

the first four abdominal vertebrae, with the exception of Grammicolepis brackiusculus, in which slender ossified ribs are

present from the third abdominal vertebra posteriorly (as

shown long ago by Shufeldt, 1888).

The term principal caudal-fin rays is used in the traditional

sense of relatively long, articulated (cross-striated) rays, which

nearly always have a single uppermost and a single lowermost

unbranched ray and branched rays between them, in contrast

to the procurrent rays, which are shorter and more spine-like

and are usually nonarticulated in acanthomorph fishes. The

single exception is Parazen pacificus, in which there are two

long, articulated, but unbranched rays above and below the

branched rays.

Both Capros and Antigonia are sometimes said to have 12

principal rays, but based on the usual definition of principal

rays being the branched rays plus the single uppermost unbranched ray and the single lowermost unbranched ray, Capros

has 14 principal rays (12 branched+ 2 unbranched) and Antigonia has 12(10 branched+ 2 unbranched).

We agree with Johnson and Patterson (1993) in their refutation of the persistent legend that some zeiforms have eight

branchiostegal rays, like most beryciforms and stephanoberyciforms, rather than seven as in most perciforms. All specimens

of zeiforms examined herein have 3 + 4 = 7 branchiostegals,

with the single exception of Macrurocyttus acanthopodus, in

which five of seven specimens have 2 + 4 = 6 branchiostegals

and two specimens have 3 + 4 (therefore, it is coded with the

majority condition of six rays).

We characterize Bremer branch support values (see "Analytical Protocols") as follows: 1 =weak, 2 = moderate, 3 = good,

4 = strong, and 5 or higher = very strong.

The term "consistent" synapomorphy is used to denote those

character states that are unambiguous, non-reversed, and nonparallel (i.e., exhibit no homoplasy) within both the 20 zeiform

taxa and the 10 outgroup taxa. As is always the case, the use of

the terms synapomorphy, apomorphy, and plesiomorphy and

our statements regarding homoplasy indicate only relative relationships. The terms are restricted to the relationships between

the taxa represented in the analyses and pertain only to those

taxa being considered for the clade under discussion.

In addition to the species examined among our outgroups,

we have consulted Zehren (1979) and Kotlyar (1996) for information on beryciforms and Tyler (1980) for tetraodontiforms;

our osteological terminology follows those works, except for a

few recently changed usages (e.g., supraneural instead of predorsal, following Mabee, 1988; rib instead of pleural rib, and

epineural instead of epipleural, following Patterson and

Johnson, 1995, and Johnson and Patterson, 2001, in contrast to

Gemballa and Britz, 1998). For the systematics of the genera

traditionally assigned to the Zeidae, we have consulted the excellent study by Bray (1983).

ZEIFORM PROCURRENT CAUDAL-FIN RAYS.—In all extant

zeiforms the principal caudal-fin rays are preceded by one to

eight procurrent rays, except in Macrurocyttus acanthopodus,

in which there are no procurrent rays (procurrent rays are also

absent in the Upper Cretaceous Cretazeus and the upper Paleocene Protozeus, but a single procurrent may be present in the

upper Paleocene Archaeozeus). The absence of procurrent rays

in the numerous specimens of Macrurocyttus, all of which are

45 mm SL or smaller (and in Cretazeus, 15-53 mm SL, and the

very small specimens of Protozeus, 9-11 mm SL) should be

considered a derived feature and not a developmental condition

because the full complement of procurrent and principal rays in

extant zeiforms is usually formed at very small sizes. We have

found the procurrent rays fully developed in postlarval or early

juvenile specimens for all of the several species of zeiforms for

which we have available specimens even smaller than those of

any of the above-mentioned species that lack procurrent rays.

For example, in the well-ossified skeleton of a cleared and

stained 8.9 mm SL specimen of Zenion hololepis (see "Material"; the caudal skeleton of this specimen is illustrated in

Johnson and Patterson, 1993:597, fig. 21), the full adult complement of 13 principal rays, three procurrent rays above the

principals, and two below the principals is present. We have

also examined a large series of alcohol-preserved postlarval

and early juvenile specimens of Z. hololepis from the collections of the Museum of Comparative Zoology (MCZ). Harvard

University. This series (MCZ 41617, 60752, 85102-85112.

87021) has 15 specimens of 6.4-20.2 mm SL, with four of

them (6.4-8.3 mm) even smaller than the smallest specimen of

even Protozeus, and all of these small specimens of Z. hololepis have the full adult complement of 2 or 3 procurrent rays
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

above and below and 13 principal rays. Likewise, a series of

six alcohol-preserved specimens of Xenolepidichthys dalgleishi of 8.4-18.7 mm SL (MCZ 57867, 57869, 57871, 57872,

84958, 84959) and a single cleared and stained specimen of

-8.9 mm SL (see "Material") all have the full adult complement of one procurrent ray above and below and 15 principal

rays, and two specimens of Parazen pacijicus of 12.5-18.7 mm

SL (MCZ 85099, 85100) have the full adult complement of 7

or 8 procurrent rays above and below and 13 principal rays.

The only small specimen of a zeiform fish we have examined

in which the caudal fin is not fully formed is a cleared and

stained 7.0 mm SL specimen of Zenopsis sp. (see "Material;"

this specimen is mentioned by Johnson and Patterson,

1993:596, as Zeus sp., 10 mm SL). The entire skeleton is unossified, and the caudal skeleton is not yet fully formed even in

cartilage; the caudal fin has five rays in the upper lobe and

seven in the lower lobe. Adults of the several species of Zenopsis have one procurrent and six principal rays above and one

procurrent and seven principal rays below; therefore, the 7.0

mm specimen of Zenopsis sp. has not yet developed two of the

uppermost rays and one of the lowermost rays.

ZEID PELVIC FIN.—The first element in the pelvic fin of

Zeus and Zenopsis has been interpreted variously as a spine

(especially in Zeus) or a ray (usually in Zenopsis). Based on

adult morphology, we consider the first element in the pelvic

fin of Zeus to be a spine because it is divided basally into two

halves for only a short distance; otherwise it is a single piece,

unsegmented and unbranched, although this spine is less robust

and more flexible than that in many other zeiforms with an unquestioned pelvic-fin spine. The pelvic-fin spine in Zeus is followed by six typical rays in Z. capensis and usually by seven in

Z. faber (sometimes only six rays in the latter). Also based on

adult morphology, we consider the first element in the pelvic

fin of Zenopsis to be a ray because it is divided into two halves

throughout its length, even though it is unsegmented and unbranched; this element is followed by five other more typical

rays, which are segmented and, with increasing specimen size,

branched. The single developmental stage we have examined

of any of the species of these two genera is a 7 mm SL larval

stage of Zenopsis sp. (Dana Collection 3915 II), and it is insufficient to settle the issue. Irrespective of its developmental history, the first element in the pelvic fin of Zenopsis is significantly different from that of Zeus; therefore, we code it

differently between the two taxa.

It is clear from the above that we do not know the true homology of the spinous compared with the ray-like first pelvicfin element in Zeus and Zenopsis; this may eventually be determined in relation to the reduction of the free pelvic radials between the bases of the rays (see Johnson, 1992; Stiassny and

Moore, 1992; Johnson and Patterson, 1993). We note, however,

that the first element of the zeiform pelvic fin, whether solid or

divided into two halves, segmented or unsegmented, is most

parsimoniously interpreted as a spine or a transformed spine.

This interpretation involves a single step, a change in morphology from spine-like to ray-like. The alternative involves at least

four steps: loss of the spine, loss of the arrector muscles associated with the spine, promotion of the first pelvic-fin ray to the

position of the spine, and recreation of the arrector muscles to

serve the "new" leading element of the pelvic fin. Arrectui

muscles in teleosts are always associated (in those taxa for

which information is available) with the leading element of the

pelvic fin where this structure is present and unreduced.

CODING CONVENTIONS.— Meristic and other characters exhibiting variability are used under the following conventions.

When one particular count or state is far more common than

any other for all of the specimens examined of a species, that

count or state is the one used in the analyses. When two or

more counts or states are about equally common among the

specimens examined for a species, the character is considered

to be polymorphic (and marked "p" in the data-set matrix (Table 1) unless the variability is confined to a single character

state having a range of values, e.g., total number of vertebrae).

In the table of meristic features (Table 2), the most common

count for a character, if there is one, is underlined. When the

materials of a particular species encompass a broad range of

sizes, statements about morphological features are based on the

larger specimens available (which, for the cleared and stained

materials, are often large juveniles or young adults), except

when significant ontogenetic changes are evident in a feature

(e.g, size of the beryciform foramen and suturing between the

ceratohyal and epihyal) and so recorded for the materials examined. Features of the few cleared and stained larval and postlarval stages examined (for Morone, Zenion, Xenolepidichthys,

and Zenopsis) are mentioned only where appropriate (e.g., development of procurrent caudal-fin rays).

ANALYTICAL PROTOCOLS.—This study primarily investigates the relationships of zeiform taxa with one another, and

secondarily the putative sister-group relationship between zeiforms and tetraodontiforms. Therefore, the morphological features selected for study are mostly those that were found to differ between various zeiform taxa and between zeiforms and a

representative of a phylogenetically basal clade of tetraodontiforms. We have not included many of the features (such as in

Stiassny and Moore, 1992, and Johnson and Patterson, 1993,

and contained references) that might clarify relationships between the lower acanthomorph outgroups used herein and zeiforms. Thus, our lack of resolution in the relationships of zeiforms and the outgroups in some of our analyses is not

surprising.

In rooting the trees, 10 potential outgroups were considered,

because a well-corroborated sister group for zeiforms has not

been established. Although tetraodontiforms and caproids

have been most frequently cited in recent years as potential

and sequential sister groups for zeiforms, the evidence for this

is equivocal (particularly for caproids, for which, additionally,

monophyly has yet to be convincingly demonstrated). All 10

potential outgroups gave the same branching sequence within

zeiforms when selected as the single outgroup. Melamphaes
NUMBER 618

was selected to root our analyses because it is generally regarded as a member of a group placed near the base of the

acanthomorph tree. Thus, even though Melamphaes is in certain ways an apomorphic basal acanthomorph, we believe it is

a more objective choice for rooting the trees than a taxon that

might be either a questionable sister group of zeiforms or one

derived either earlier or later than the common ancestor of zeiforms. Moreovci, even representatives of the more basal

clades of tetraodontiforms are so osteologically apomorphic as

to be of somewhat limited value in the polarization of zeifonn

characters.

We recognize the limitations in using a single species to represent a large higher category (e.g., Morone representing percoids, and Parahollardia representing tetraodontiforms) and

that our hypothesis-testing is of the relationships only of the included taxa (or, rather, of a limited number of specimens that

we presume adequately represent a species).

Data were gathered on 124 morphological features for 20

zeiform and 10 outgroup taxa. Of these features, 17 were discarded from further consideration early in the study for a variety of reasons; they were either too intraspecifically variable or

too subjective to determine consistently between taxa, or they

over-emphasized certain repetitive configurations, such as the

number of pterygiophores between neural and haemal spines of

long sequences of successive vertebrae.

Of the remaining 107 putatively informative characters examined, the number was further reduced to 103 (Table 1) by removing four characters that were autapomorphic and thus phylogenetically uninformative: presence/absence of nasals

(absent only in Parahollardia lineata and all other tetraodontiforms), presence/absence of teeth on the premaxilla (absent

only in Oreosoma atlanticum), length of the gill opening (reduced to a short slit only in Parahollardia lineata and all other

tetraodontiforms), and number of unbranched principal caudalfin rays (increased from two to four only in Parazen pacificus).

A second, more restricted, data set was produced by removing 11 meristic characters. Serially repeated, or "number line,"

characters, such as meristic features, are difficult to interpret in

phylogenetic analyses (Murphy and Doyle, 1998). In addition,

meristic characters can be problematic because of the effects of

varying developmental condition (see "Meristic CharacterState Evolution," below).

The 11 meristic characters (93 through 103) are as follows:

total number of vertebrae, number of abdominal vertebrae,

number of vertebrae in the caudal peduncle, number of principal caudal-fin rays, number of procurrent caudal-fin rays, number of dorsal-fin spines, number of vacant interneural spaces,

number of anal-fin spines, number of pectoral-fin rays, total

number of pelvic-fin elements, and number of branchiostegal

rays.

The data for the 20 zeiforms and 10 outgroups were analyzed

both as the full data set of 103 characters and as a reduced data

set of 92 characters. All of these characters and their states are

defined at the beginning of the "Results" section.

Most multistate characters were coded and ordered as linear

transformation series; the exceptions to this are noted in the

character-state descriptions. The data set was also examined

with all characters as unordered transformation series. The data

were analyzed using Hennig86 (Farris, 1988). Because of the

large number of taxa in the data set, implicit enumeration ("ie")

was not appropriate; therefore, the Wagner trees were created

by multiple passes through the data matrices ("mh"), and then

branch breaking and swapping ("bb*") was applied to find the

most parsimonious tree(s). Before analysis in Hennig86, the order of taxa in all data matrices was randomized using the Joyride program in Random Cladistics (Siddal, 1997).

In all four analyses (ordered and unordered, both with and

without 11 meristic characters), Melamphaes served as the primary outgroup and rooted the trees. The polymorphic data

were analyzed using mutation coding as it applies to morphological characters (Murphy, 1993). The polymorphic cells were

coded as unknown ("?") in the initial analyses, then the characters were optimized onto the resulting tree(s), and the polymorphic data were reassessed. Each polymorphic cell was either recoded, representing the novel evolutionary state (i.e., the

mutation), or left unchanged, if the transformation was unresolved or ambiguous. The entire data set was then reanalyzed

and the remaining polymorphic cells were reassessed in the

same manner. This process was repeated until there were no

changes in the data set.

Characters were optimized using the accelerated transformation (ACCTRAN) option in CLADOS (Nixon, 1992), in which

reversals are favored over parallelisms for homoplastic characters. Ambiguous character optimizations (i.e., ACCTRAN

maps the character differently than in delayed transformation,

DELTRAN) are noted in the section "Phylogenetic Results."

The support for each clade, as branch-support values (Bremer,

1988, 1994), were evaluated using the "branch support" option

in NONA (Goloboff, 1993).

Ten thousand trees were generated that were a maximum of

15 steps longer than the most-parsimonious tree; consensus

trees were derived from these to determine branch support.

These represent the number of extra steps needed to be added

to the most-parsimonious tree that will collapse that branch in

the strict consensus of all trees of that length and shorter. These

values are considered to be good indicators of ranked nodal

support (Doyle, 1998).

TREE CHARACTERISTICS.—Using the full data set of 103

characters, the ordered analysis resulted in six trees of 581

steps (CI=40, RI=65); the strict consensus tree is shown in

Figure 1. The unordered analysis resulted in a single most-parsimonious tree of 482 steps (CI=47, Rl = 64), which is shown

in Figure 2.

Using the reduced data set of 92 characters, the ordered analysis resulted in 11 trees of 386 steps (Cl = 43, Rl = 66); the

strict consensus tree is shown in Figure 3. The unordered analysis resulted in 26 trees of 354 steps (CI = 46, Rl = 68); the

strict consensus tree is shown in Figure 4.
SMI1 HSONIAN CONTRIBUTIONS TO ZOOLOGY

L=581

CI=40

RI=65

Melamphaes suborbitalis

Anoplogaster cornuta

Hoplostethus mediterraneus

Centroberyx affinis

Morone americana

Plectrypops lima

Sargocentron punctatissimum

Capros aper

• Antigonia capros

• Parahollardia lineata

• Cyttus traversi

• Cyttus australis

• Cyttus novaezelandiae

• Zenion hololepis

• Capromimus abbreviatus

• Cyttomimus stelgis

• Parazen pacificus

• Cyttopsis roseus

• Stethopristes eos

• Pseudocyttus maculatus

• Allocyttus verrucosus

• Neocyttus rhomboidalis

• Allocyttus niger

• Oreosoma atlanticum

• Macrurocyttus acanthopodus

• Xenolepidichthys dalgleishi

• Grammicolepis brachiusculus

• Zeus faber

• Zenopsis conchifer

• Zenopsis nebulosus

FIGURE 1.—Strict consensus of the six most-parsimonious trees resulting from an ordered analysis of the complete data set of 103 characters. Branch support indices (Bremer, 1994) are shown above all relevant branches.

The tree is rooted at Melamphaes.
NUMBER 618

L=482

CI=47

RI=64

Melamphaes suborbitalis

Anoplogaster cornuta

Plectrypops lima

Sargocentron punctatissimum

Hoplostethus mediterraneus

Centroberyx affinis

Morone americana

Capros aper

Antigonia capros

Parahollardia lineata

Cyttus novaezelandiae

Cyttus australis

Cyttus traversi

Pseudocyttus maculatus

Neocyttus rhomboidalis

Allocyttus verrucosus

Allocyttus niger

Oreosoma atlanticum

Parazen pacificus

Cyttopsis roseus

Stethopristes eos

Zenion hololepis

Capromimus abbreviatus

Cyttomimus stelgis

Macrurocyttus acanthopodus

Xenolepidichthys dalgleishi

Grammicolepis brachiusculus

Zeus faber

Zenopsis conchifer

Zenopsis nebulosus

FIGURE 2.—Single most-parsimonious tree resulting from an unordered analysis of the complete data set of 103

characters. Branch support indices (Bremer, 1994) are shown above all relevant branches. The tree is rooted at

Melamphaes.
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

L=386

CI=43

RI=66

— Melamphaes suborbitalis

— Morone americana

— Sargocentron punctatissimum

— Plectrypops lima

— Centroberyx affinis

— Hoplostethus mediterraneus

— Anoplogaster cornuta

— Capros aper

— Antigonia capros

Parahollardia lineata

Cyttus australis

Cyttus novaezelandiae

Cyttus traversi

Pseudocyttus maculatus

Oreosoma atlanticum

Neocyttus rhomboidalis

Allocyttus verrucosus

Allocyttus niger

Parazen pacificus

Cyttopsis roseus

— Stethopristes eos

— Zenion hololepis

— Capromimus abbreviatus

— Cyttomimus stelgis

— Macrurocyttus acanthopodus

— Xenolepidichthys dalgleishi

— Grammicolepis brachiusculus

— Zeus faber

— Zenopsis conchifer

Zenopsis nebulosus

FIGURE 3.—Strict consensus of the 11 most-parsimonious trees resulting from an ordered analysis of the reduced

data set of 92 characters. Branch support indices (Bremer, 1994) are shown above all relevant branches. The tree

is rooted at Melamphaes.
NUMBER 618 11

A

B

C

D

t

2

F

G

H

1

J

K

10

L

L=354

CI-46

RI=68

7

M

1

R

N

1

0

2

S

2

A

1

2

B

1

1

G

1

P

Q

T

2

U

C

3

D'

2

H'

1

M'

V

W

1

X

Y

Z

E'

F"

r

2

J'

4

N'

12

S'

K'

L

1

0'

8

P'

T

2

U'

Q'

R'

V

w

Melamphaes suborbitalis

Anoplogaster cornuta

Hoplostethus mediterraneus

Centroberyx affinis

Morone americana

Plectrypops lima

Sargocentron punctatissimum

Capros aper

Antigonia capros

Parahollardia lineata

Cyttus novaezelandiae

Cyttus australis

Cyttus traversi

Pseudocyttus maculatus

Oreosoma atlanticum

Neocyttus rhomboidalis

Allocyttus verrucosus

Allocyttus niger

Parazen pacificus

Cyttopsis roseus

Stethopristes eos

Zenion hololepis

Capromimus abbreviatus

Cyttomimus stelgis

Macrurocyttus acanthopodus

Xenolepidichthys dalgleishi

Grammicolepis brachiusculus

Zeus faber

Zenopsis conchifer

Zenopsis nebulosus

FIGURE 4.—Strict consensus of the 26 most-parsimonious trees resulting from an unordered analysis of the

reduced data set of 92 characters. Branch support indices (Bremer. 1994) are shown above all relevant

branches. Letters below branches refer to the list of character-state transformations in the Appendix. The tree is

rooted at Melamphaes.
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

In Figures 1-4, branch-support values (Bremer, 1988, 1994)

are indicated above all nonterminal branches, and in Figure 4

the letters below the branches represent the character transformations, which are given in the Appendix.

The character-state distributions generated by CLADOS for

the reduced data set (Figures 3, 4) are considered here to be

more rational than those for the full data set because of the

problematic nature of many meristic characters, especially in

zeiforms (see "Meristic Character-State Evolution"). Additionally, for the reduced data set, the unordered tree (Figure 4) is

better justified than the ordered tree (Figure 3), because we

have no unequivocal ontogenetic or other data that might allow

us to order individual transformation series. Not surprisingly,

our attempts at ordering were largely unsuccessful and resulted

in trees that were longer than that given in Figure 4. Extra steps

were especially apparent for multistate characters. The unordered, reduced-data-set tree (Figure 4) is 32 steps shorter than

the tree generated with the ordered, reduced data set (Figure 3).

The 11 meristic characters, although not used in constructing

the trees, are included in the character distributions used to

evaluate nodal support (Figure 5), but they are not included in

branch-support calculations.

The use of consensus trees in reconstructing character evolution is somewhat problematic (Maddison, 1989), because the

length of the consensus tree may be longer than the most-parsimonious trees. The consensus tree of the unordered, reduced

data set (Figure 4) is 39 steps longer than any of the 26 mostparsimonious trees. These extra steps result from the need to

interpret the characters that map on the large, basal polytomy

and on the single trichotomy of the consensus tree. The large

polytomy occurs in the outgroups, so the extra steps added here

do not affect our conclusions about zeiforms. Characters 36

and 58, however, both require additional steps on the consensus

tree within zeiforms because of the trichotomy involving Oreosoma, Neocyttus, and Allocyttus (Figure 4). By analyzing the

evolution of these characters on all 26 equally parsimonious

trees, we find that they reconstruct different distributions, and,

thus, they should be used with caution to support this group.

The degree of congruence of the 11 meristic characters removed from the analyses was evaluated by optimizing them

onto the consensus tree resulting from the unordered analysis

of the reduced data set (Figure 4) (see "Meristic CharacterState Evolution" and Figure 5). This resulted in one meristic

character being congruent with the phylogeny, four others remaining questionable, and six being rejected for future use in

zeiforms.

Statements about monophyly of the various taxa are based

only on the analysis of the unordered, reduced data set (Figure

4) for the species examined (see "Material").

MERISTIC CHARACTER-STATE EVOLUTION.—Many of the

serially repeated structures of zeiforms have significant variability within and between geographic populations of a species,

especially for the soft dorsal- and anal-fin rays. This variability

is from a combination of genetic factors and environmental

factors, such as temperature and oxygen levels during early

life-history stages (Lindsay, 1988). acting on the genomes

within a population, and from differences in these factors between populations. It is almost impossible on the basis of the

highly limited samples of zeiforms available from museum

specimens alone (collected at diverse places and at different

times, and rarely a statistically significant sample of any one

population at one time), in the absence of information on developmental conditions and population dynamics, to separate

how much of the observed meristic variation is phenotypic versus genetic.

The soft dorsal- and anal-fin rays are sufficiently variable intraspecifically, including within the few large samples available of the same population of some of the zeiform taxa examined, that no attempt was made to use interspecific differences

in the counts of soft fin-rays.

Moreover, among zeiforms even some of the usually more

conservative and consistent structures, such as fin spines, are

relatively more variable, and there is more lability between

spines and rays than in most other acanthomorphs. For example, the first element in the pelvic fin of two closely related

genera (Zeus and Zenopsis) is spine-like in one and ray-like in

the other; the anal-fin spines vary between four and five in

Zeus faber and between two and three or between three and

four in several species of oreosomatids; the pelvic-fin rays may

vary by one element in several species of several families; the

relatively low number of abdominal vertebrae varies by one

and occasionally two units in several taxa of oreosomatids and

zeids (but partially because of difficulty in definition; see "Terminology"); and even the relatively conservative principal caudal-fin ray count is variable in one species (Macrurocyttus

acanthopodus).

We realize that the more qualitative features, such as the

shapes and sizes of bones, that are the basis for other characters

used in this study also exhibit variability, but this is usually of a

lesser degree than with many of the meristic characters, and the

range of conditions is contained within our definition of the

character. We recognize that the variability in qualitative characters is more difficult and subjective to determine and describe than it is for easily quantified meristic features, and that

this accounts for variability in qualitative features usually not

being taken into consideration. But because many of the meristic features are especially variable in zeiforms, and we cannot

distinguish between the phenotypic and genetic components of

that variability, their value in a phylogenetic analysis is perhaps

questionable, and shared conditions may not be due exclusively

to homology.

Therefore, we think it is reasonable to exclude most of the

fin and vertebral meristic characters from the analyses and then

to optimize them onto the cladogram to test congruence. By

comparing the distribution of each of these characters with respect to the phylogeny of the group, we can either accept or reject this a priori assumption. If the character is congruent (high

ci and ri) with the phylogeny, then the observed variation is
NUMBER618 13

93-2,95-8,974,98-0, 99-0,100-1,102-1

93-1, 94-4,95-7, 97-7, 98-0, 99-0,100-0,102-2

98-2, 99-0,101-4,102-2

94-1,97-5,99-0,100-4,101-1,102-1

94-3, 97-9.103-1

93-1,102-1

95-3,1004,101-3

94-1,95-1,96-2,98-5,103-2

94-1, 95-0,964, 984,101-1,103-2

94-0,95-2, 964, 97-0, 98-2, 99-2,100-0,101-1,102-6,103-2

93-3, 96-3, 984,

100-2,102-2,103-1

L=354

CI=46

RI=68

95-2,

101-0

95-3,

97-3,

98-3

93-8, 94-5,

99-8,1014

93-8,97-3,98-5,99-2,101-3

98-1,99-7,102-3

100-1

93-7,

97-2

93-6, 954,100-3

97-3, 99-9

93-6, 944, 98-2, 99-7

954,100-3

934, 94-3, 954, 984

99-3,

101-1,

102-0

974, 98-2

99-5

99-2

93-1,95-2,97-2,98-2

954, 974

93-2, 984.99-1

93-5, 95-2,

97-1,101-1

95-7

944,95-1,97-0,100-0,

102-5,103-2

93-9,96-1,

994

93-6,94-1,

95-5,98-2

101-2

94-5. 98-6.

1004.102-3

93-3.

102-1

100-3,

101-0

94-6,

95-1

• Melamphaes suborbitalis

• Anoplogaster cornuta

Hoplostethus mediterraneus

Centroberyx affinis

Morone americana

Plectrypops lima

Sargocentron punctatissimum

Capros aper

Antigonia capros

Parahollardia lineata

Cyttus novaezelandiae

Cyttus australis

Cyttus traversi

Pseudocyttus maculatus

Oreosoma atlanticum

Neocyttus rhomboidalis

Allocyttus verrucosus

Allocyttus niger

Parazen pacificus

Cyttopsis roseus

Stethopristes eos

Zenion hololepis

Capromimus abbreviatus

Cyttomimus stelgis

Macrurocyttus acanthopodus

Xenolepidichthys dalgleishi

• Grammicolepis brachiusculus

• Zeus faber

Zenopsis conchifer

• Zenopsis nebulosus

FIGURE 5. Optimization of the 11 meristic characters (characters 93-103) onto the strict consensus tree of the

unordered reduced data set of 92 characters in FIGURE 4, to test for congruence with the phylogenv. The tree is

rooted at Melamphaes.
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

most likely a function of genealogy and the shared states are

homologous. If the character is not congruent (low ci and ri),

then the variation may be a function of something other than

phylogeny, and the shared states may be convergent.

The 11 meristic characters (characters 93 through 103) that

were excluded from the data set were optimized onto the cladogram (Figure 5) to determine their congruence with our hypothesis of genealogy (Figure 4). Each character was optimized

separately, and the ci and ri are noted for each character so that

conclusions can be drawn about their congruence with the phylogeny. The values of these indices are compared with those of

the unordered consensus tree (CI = 46, RI = 68), with "good"

and "poor" defined as higher or lower than those values, respectively. This test of congruence will only determine to what

degree the observed variation reflects phylogeny, and, to that

extent, the usefulness of a character in a phylogenetic analysis.

Many of these characters are regularly used in systematic characterizations, and their great utility for that purpose is not being

questioned.

Character 93, total number of vertebrae (ci = 56, ri = 53, 16

steps for 9 states). This character exhibits a moderate amount

of homoplasy, as shown by the seven extra steps needed to map

its distribution. The synapomorphy support, ri, is also moderate. Therefore, the usefulness of this character in this phylogenetic analysis remains uncertain.

Character 94, number of abdominal vertebrae (ci = 46, ri =

22, 13 steps for 6 states). This character exhibits a moderate

amount of homoplasy, six extra steps, and poor synapomorphy

support. The poor synapomorphic support results from mapping character-state transitions on many terminal branches

(e.g., Centroberyx and Morone).

Character 95, number of caudal-peduncle vertebrae (ci = 38,

ri = 21, 18 steps for 8 states). This character also exhibits a

large amount of homoplasy, six extra steps, and poor synapomorphy support.

Character 96, number of principal caudal-fin rays (ci = 80,

ri = 87, 5 steps for 4 states). This character exhibits little homoplasy; for this group of fishes the number of caudal-fin rays

is a good indicator of phylogeny.

Character 97, number ofprocurrent caudal-fin rays (ci = 50,

ri = 36, 14 steps for 7 states). This character exhibits a moderate amount of homoplasy and poor synapomorphy support.

Character 98, number of dorsal-fin spines (ci = 35, ri = 21, 17

steps for 6 states). This character exhibits a large amount of

homoplasy and little synapomorphy support.

Character 99. total number of vacant interneural spaces

(ci = 53, ri = 46, 12 steps for 9 states). This character exhibits

a moderate amount of homoplasy and moderate synapomorphy

support.

Character 100, number of anal-fin spines (ci = 33, ri = 46, 12

steps for 4 states). This character exhibits a large amount of

homoplasy and poor synapomorphy support.

Character 101, number of pectoral-fin rays (ci = 33, ri=46,

12 steps for 4 states). This character exhibits a large amount

of homoplasy and moderate synapomorphy support.

Character 102, total number of pelvic-fin elements (ci = 41,

ri = 30, 12 steps for 5 states). This character exhibits a moderate amount of homoplasy and poor synapomorphy support.

Character 103, number of branchiostegal rays (ci = 33, ri =

50, 6 steps for 2 states). This character exhibits a large

amount of homoplasy and moderate synapomorphy support.

Of these, only character 96, number of principal caudal-fin

rays, is congruent with our hypothesis of phylogeny. This character is presumably affected by variation in developmental

conditions acting on differential genomes between semi-isolated populations, but this variation does not mask the phylogenetic signal.

Four of the other meristic characters are of uncertain utility:

total number of vertebrae (93), number ofprocurrent caudal-fin

rays (97), total number of vacant interneural spaces (99), and

number of branchiostegal rays (103). These characters presumably are also affected by the interaction of developmental conditions on population genomes, but we cannot determine if the

variation between taxa is a result of phylogeny or some other

factor. Because we cannot determine if these shared traits are

due to convergence or homology, the value of these characters

in this phylogenetic analysis remains questionable.

The remaining six meristic characters show very little congruence with the hypothesis of phylogeny, and they have poor

synapomorphic support: number of abdominal vertebrae (94),

number of caudal-peduncle vertebrae (95), number of dorsalfin spines (98), number of anal-fin spines (100), number of

pectoral-fin rays (101), and total number of pelvic-fin elements

(102). There is little evidence that these character-state distributions reflect the phylogeny of these fishes.

ILLUSTRATIONS.—Instead of detailed descriptions of the

comparative osteological features and character states of zeiforms, we provide illustrations of representatives of each of the

six families recognized herein. For each family, an illustration

of the lateral view of the entire skeleton of a representative species is given, as well as detailed figures of phylogenetically informative features of its skeleton or skeletal features in comparison with those of other species of the same or subsequently

treated families. These detailed illustrations are labeled with the

character number separated by a dash from the state number; in

the figure legends, some of the character numbers are given

only abbreviated descriptions. To preserve the unimpeded overall configuration of the bony parts in the lateral views of the entire skeleton of representative species, pointer lines are not used

for the mostly self-evident character states given in the legends.

Illustrations of entire skeletons were prepared from outlines

based on projections from 35 mm photographic slides of intact

whole skeletons (all cleared and stained except one dry skeleton

and one radiograph) and refined by microscopic examination of

the specimen; all detailed illustrations were drawn using a camera lucida microscope attachment.
NUMBER 618 15

All of the osteological illustrations are arranged at the back

of this publication in phylogenetic sequence: Cyttidae, Oreosomatidae, Parazenidae, Zeniontidae, Grammicolepididae, and

Zeidae.

Scale bars in the illustrations are in mm. A single scale bar

for multiple figures applies to all of the parts.

Lateral views of details of the skeleton are of the left-side

parts unless otherwise stated.

Gill rakers are not shown in the branchial arch illustrations.

Cartilages and dense fibrous tissue in the branchial arches

and elsewhere in the skeleton are not consistently illustrated;

when shown, these areas are outlined and are either open, without stippling, or have mechanical stippling or parallel or wavy

lines.

Material

Specimens examined are listed for each species in the following order: catalog number, number of specimens, standard

length (mm), general geographic region of specimen collection,

and type of preparation, with CS = cleared and stained (usually

counter stained for cartilage), D = dry skeletal preparation, and

R = radiograph. CS specimens are listed first, followed by D

and R specimens; within each of these categories, specimens

are listed alphabetically by institutional abbreviation, which

follow Leviton et al. (1985). The zeiform species are listed alphabetically within the new family arrangements proposed

herein, followed by the outgroup species. Author names for

species are given here rather than throughout the text.

ZEIFORMES

CYTTIDAE

Cyttus australis (Richardson). AMS 1.32386001, 1 (60), Australia, CS.

AMNH 91908, 1 (302), Australia, D; AMNH 91910, 1 (233), Australia,

D; AMNH 91932, 1 (250), Australia, D. USNM 177052, 2 (168, 183),

Australia, R.

Cyttus novaezelandiae (Arthur). AMNH 95882. 1 (155), Australia, D;

AMNH 95891, 1 (162), Australia, D. USNM 320593, 1 (263), New

Zealand, R.

Cyttus traversi Hutton. AMNH 92029, 1 (316), Australia, D; AMNH 92052,

1 (215), Australia, D; AMNH 99777, 1 (455), Australia, D; AMNH

99778, 1 (445), Australia, D. USNM 308020, 1 (105), Walters Shoal, Indian Ocean, R; USNM 320583, I (263), New Zealand, R.

OREOSOMAT1DAE

Allocyttus folletti Myers. USNM 306419, 1 (250), Alaska, R. Data from radiograph insufficient to include species in data matrix and analyses.

Allocyttus niger James, Inada, and Nakamura. AMNH 98405, 1 (315), Australia, D; AMNH 98407, 1 (315), Australia, D; AMNH 98408, 1 (350), Australia, D; AMNH 98629, 1 (320), Australia, D; AMNH 98630, 1 (325),

Australia, D; AMNH 98634, 1 (345), Australia, D.

Allocyttus verrucosus (Gilchrist). AMS I.18712OO2, 1 (83), Australia, CS;

AMS I.2OO68O31, 2 (89, 90), Australia, CS. AMNH 95091, 1 (252),

Australia, D; AMNH 95651, 1 (280), Australia. D; AMNH 95652, I

(260), Australia, D; AMNH 95653, 1 (265), Australia, D. USNM

216741, 5 (272-300), Madagascar, R; USNM 307599, 3 (174-182),

Madagascar, R; USNM 320678, 1 (250), Madagascar, R; USNM 320709,

2 (275, 279), Madagascar, R.

Neocyttus rhomboidalis Gilchrist. AMS 1.20099029, I (87), Australia, CS;

AMS 1.21812002. 1 (81). Australia, CS. AMNH 91743, 1 (295), Australia, D; AMNH 91746, 1 (232), Australia, D; AMNH 93612, 1 (243), Australia, D; AMNH 93613, 1 (255), Australia, D; AMNH 95080, 1 (248),

Australia, D. USNM 320680, 1 (187), Australia, R.

Oreosoma atlanticum Cuvier. AMS 1.21370034, 1 (32), Australia, CS; AMS

1.25127008, 1 (133), Australia,CS; AMS 1.25933013, 1 (109), Australia,

CS. AMNH 99737, 1(119), Australia, D.

Pseudocyttus maculatus Gilchrist. AMNH 95075, 1 (195), Australia, D;

AMNH 95088, 1 (255), Australia, D.

PARAZENIDAE

Cyttopsis roseus (Lowe). AMNH 29460, 2 (74, 75), Nicaragua, CS; AMS

1.19840010, 1 (46), Australia, CS; CAS 38408, 1 (81), Gabon, CS;

USNM 187873, 1 (72), Costa Rica, CS; USNM 304460, 2 (62, 67), Nicaragua, CS. USNM 50562, 1 (130), Japan, R (holotype of synonymous

Cyttopsis itea Jordan and Fowler); USNM 102129, 1 (182), Puerto Rico,

R; USNM 204892, 2 (385, 412), Cuba, R; USNM 227636, 1 (92), locality unknown, R; USNM 227936, 1 (193), Colombia, R; USNM 240051,

1 (95), Bermuda, R.

Parazen pacificus Kamohara. AMNH 29459, 1 (105), Florida, CS; CAS

38404, 1 (96), Nicaragua, CS; USNM 187077, 1 (104), Puerto Rico, CS;

USNM 187892, 1(110), Panama, CS. USNM 157874, 1 (93), Cuba, R;

USNM 187077, 2 (96, 97), Puerto Rico, R; USNM 187892, 1 (128), Panama, R; USNM 927761, 1 (174), Columbia, R. Osteology well described

byKotlyar(2001).

Stethopristes eos Gilbert. USNM 226570, 2 (91, 97), Chile, CS. USNM

51626, 1 (107), Hawaii, R (holotype); USNM 226570, 2 (102, 116),

Chile, R; USNM 334054, 2 (108, 110), Chile, R.

ZENIONTIDAE

Capromimus abbreviatus (Hector). AMS I.23650OO1, 1 (61), New Zealand,

CS. USNM 334056, 1 (68), New Zealand, R.

Cyttomimus stelgis Gilbert. USNM 306156, 1 (76), Somalia, CS. AMS

1.20651018, 2 (73, 81), Australia, R; USNM 51622, 1 (76), Hawaii, R

(holotype); USNM 226569, 2 (46, 67), Chile, R; USNM 334055, 3

(55-66), Chile, R.

Zenion hololepis (Goode and Bean). CAS 38409, 1 (70), Bahamas, CS: CAS

76856, 2 (70, 73), no locality, CS; Dana Collection (Copenhagen), station 3744 II, uncataloged, 1 (9), western North Atlantic, CS (well-ossified larval stage); USNM 187864, 3 (50-73), Panama. CS; USNM

307305, 2 (45, 48), Madagascar, CS. USNM 138338, 9 (52-69), Philippines, R; USNM 138339, 1 (54), Philippines, R; USNM 138340. 1 (45).

Philippines, R; USNM 138341, 1 (32), Philippines, R; USNM 327842, 4

(60-70), Venezuela, R; USNM 334057, 5 (47-58), Chile, R.

Zenion leptolepis (Gilchrist and von Bonde). USNM 320058, 10 (43-49).

Liberia, R; USNM 320059, 3 (83-90), Liberia, R. Data from radiographs

insufficient to include species in data matrix and analyses.

GRAMMICOLEPIDIDAE

Grammicolepis brachiusculus Poey. AMNH 99621, 1 (285), Australia. D.

USNM 84098, I (226), Hawaii, R (holotype of synonymous Vesposus

egregius Jordan); USNM 102129, 1 (182), Puerto Rico, R; USNM

204892, 2 (385, 412), Cuba, R; USNM 227636. 1 (92), no locality, R;

USNM 227936. 1 (193), Colombia (Atlantic), R; USNM 240051. 1 (95).

Bermuda, R. Osteology well described by Shufeldt (1888).

Macrurocyttus acanthopodus Fowler. MNHN 1994-1072, 1 (43), New Caledonia, stained, not cleared; USNM 93144, 5 neurocrania 13-14 mm long

(vomer to basioccipital) and assorted disarticulated bones from type

specimen series (in 14 glass vials), only one specimen, alcohol preserved, intact (removed and recataloged as USNM 331213, see following

entry), Philippines, stained, not cleared, all flesh absent. MNHN 1994-

1052, 2 (37, 39), New Caledonia. R; MNHN 1994-1073. 3 (37^5). New

Caledonia, R; USNM 331213, 1 (-36), Philippines, R (a type specimen,

but cataloging data and data published by Fowler (1934) conflict; impossible to determine whether it is the holotype). Rarely collected; see

Nyako and Amaoka (1996) for redescription of species.

Xenolepidichthys dalgleishi Gilchrist. AMNH 29455, 4 (70-75). Nicaragua.

CS; CAS 38403, 1 (70). Nicaragua. CS: CAS 38406. 1 (75). Colombia.

CS; MCZ 57872. I (-8-9), Caribbean. CS; USNM 320013. 1 (64). Hon-
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

duras, CS; USNM 320015, 3 (59-77), Honduras, CS; USNM 320016. 2

(59, 64), Panama, CS. USNM uncataloged, 4 (77-90), Honduras. R;

USNM 328266, 4 (69-75), Honduras, R; YPM (BOC) 517, 1 (82), Belize, R; YPM (BOC) 518, 2 (73, 81), Belize, R; YPM (BOC) 524, 1 (85),

Belize, R.

ZEIDAE

Zenopsis conchifer (Lowe). CAS 47401, 1 (54), Florida, CS; FMNH 67090,

3 (44-58), Florida, CS; USNM 117280, 2 (51, 81), Florida, CS. AMNH

4451, 1 (126), New York, D; AMNH 56447, 1 (368), New York, D;

AMNH 56833, 1 (405), New York, D.

Zenopsis nebulosus Schlegel. AMNH 92291, 1 (325), Australia, D; AMNH

95024, 1 (335), Australia, D; AMNH 95028, 1 (340), Australia, D.

Zenopsis oblongus Parin. USNM 334160, 2 (one partial skeleton, front of

head missing, -280 mm SL; one cranium from specimen of similar

size), Nazca Ridge, southeast Pacific, D. USNM 353898, 1 (347), Salay-Gomez Ridge, southeast Pacific, R. Data from radiograph and two incomplete skeletons insufficient to include species in data matrix and

analyses.

Zenopsis sp. Dana Collection (Copenhagen), station 3915 II, uncataloged, 1

(7), western North Atlantic, CS (weakly ossified larval stage, probably of

Z. conchifer).

Zeus capensis Valenciennes. USNM 330849, 7 (173-193), South Africa, R.

Data from radiographs insufficient to include species in data matrix and

analyses.

Zeus faber Linnaeus. USNM 307842, 2 (48, 67), Tunisia, CS; USNM

320014, 2 (59, 89), Tunisia, CS; USNM 320063, 1 (80), Guinea, CS.

AMNH 22707, 1 (300), Japan, D; AMNH 91448, 1 (230), Australia, D;

AMNH 95054, 1 (280), Australia, D; AMNH 95055, 1 (270), Australia,

D; USNM 176975, 1 (215), Australia, D. USNM 328597, 1 (223),

Kenya, R.

OUTGROUPS

TRIACANTHODIDAE

Parahollardia lineata (Longley). ANSP 93375, 1 (62), Florida, CS; ANSP

97637, 3 (79-84), Louisiana, CS; ANSP 100473, 1 (46), Florida, CS;

USNM 280326, 1 (121), Louisiana, CS. Osteology described in detail by

Tyler (1968, 1980).

CAPROIDAE

Antigonia capros Lowe. AMNH 21929, 1 (48), Florida, CS; AMNH 29461,

3 (31^43), Nicaragua, CS; FMNH 64378, 1 (50), Nicaragua, CS; USNM

289209, 4 (30-47), Colombia, CS; USNM 320064, 2 (38, 40), Colombia. CS. Osteology of related A. comhatia Berry and Rathjen described

by Zehren (1987). Osteology of A. rubescens (Giinther) compared with

that of chaetodontoids by Starks (1902).

Capros aper (Linnaeus). USNM 268912, 5 (40-73), Morocco, CS; USNM

289207, 3 (48-65), Tunisia, CS; USNM 320065, 2 (57, 58), Morocco,

CS. Osteology briefly described by Tortonese (1948); described in great

detail by Zehren (MS).

HOLOCENTRIDAE

Plectrypops lima Valenciennes. USNM 288860, 1 (61), Tonga, CS; USNM

334646, 2(50,52), Fiji, CS.

Sargocentron punctatissimum (Cuvier). USNM 259019, 2 (53, 61), New

Guinea, CS; USNM 269081, 2 (54, 70), Fiji, CS.

MORONIDAE

Morone americana Gill. ROM 369CS, 2 (40, 44), no locality, CS; ROM

373CS, 1 (115), Ontario. CS; ROM 1064CS, 1 (86), Ontario, CS;

USNM 109851, 3 (42-46), Virginia, CS; USNM 187300, 1 (77), Maryland, CS; USNM 322796, 30 (7-12), reared larvae, CS. ROM 2605, 1

(320), Ontario, D.

BERYCIDAE

Centroberyx affmis (Giinther). USNM 176984, I (110), Australia, CS. Osteology described in detail by Zehren (1979). Osteology treated by Kotlyar

(1996).

TRACHICHTHYIDAE

Hoplostethus mediterraneus (Cuvier and Valenciennes). USNM 306630, 1

(61), Somalia, CS; USNM 306631, 2 (50, 61), Somalia, CS. Osteology

treated by Kotlyar (1996).

ANOPLOGASTERIDAE

Anophgasler cornuta (Valenciennes). USNM 214217, 1 (38), Mississippi,

CS; USNM 240793, 1 (43), Bermuda, CS; USNM 244922, 1 (26), Bermuda, CS; VIMS uncataloged, I (93), no locality, CS. USNM 215619, 1

(103), Hawaii, R; USNM 240772, 2 (97, 124), Bermuda, R; USNM

343725, 1 (80), Florida, R. Osteology described in detail by Zehrc;,

(1979).

MELAMPHA1DAE

Melamphaes suborbitalis (Gill). USNM 218556, 1 (62), South Africa, CS;

USNM 218562, 1 (63), South Africa, CS. Osteology well described by

Kotlyar (1999).

Results

PHYLOGENY

Characters and Character States

The 103 characters and their states (coded as ordered transformation series where we deem it appropriate) are organized

and discussed according to anatomical regions. Characters for

which we are not confident about possible transformation series, and therefore treat as unordered, are indicated as such. The

distribution of the character states among the taxa analyzed is

given in Table 1.

Cranium

CHARACTER 1.—Parietal: present (0); absent (1).

CHARACTER 2.—Basisphenoid: present as a long, oblique

shaft connecting parasphenoid and prootic in front of posterior

myodome (0); present as a short shaft at front of roof of posterior myodome (1); absent (2).

CHARACTER 3.—Vomerine teeth: present (0); absent (1).

CHARACTER 4.—Parasphenoid opening into posterior myodome: absent (0); present (1).

CHARACTER 5.—Honeycomb bone sculpturing on skull,

opercles, and lachrymal-infraorbitals: absent (0); present (1).

CHARACTER 6.—Supraocular serrations on frontal: present

(0); absent (1).

CHARACTER 7.—Otolith shape and size: moderately large to

large, rounded or slightly to deeply indented on one or both

sides, or oblong with humps (0); tiny, trilobed (bow-tie

shaped) (1).

We have examined zeiform and outgroup otoliths for this

work but have not illustrated them; for illustrations of zeiform

otoliths see Stinton (1967), Nolf (1985), Nolf and Cappetta

(1988), Smale et al. (1995), and Schwarzhans (1996).

CHARACTER 8.—Size and depth of lachrymal: large, deep,

height about two to four times in the length (0); moderately

deep, height about five to seven times in the length (1); slender

(2); absent (-).
TABLE 1.—Data set for the 30 taxa in the analyses: "-"=inapplicable; "?"=unknown, i.e., cannot be determined in

our materials; "p"=polymorphic character with more than one character state common in our materials. For purposes of the analyses, the "?" and "p" categories are treated equally. Characters marked "*" were unordered in all

analyses. Polymorphic data are as follows. Hoplostethus mediterraneus p=97-6,7. Centmberyx affinis p=98-3,4.

Morone americana p=71-l,2; 72-2,3; 101-2,3. Plectrypops lima p=97-5,6. Sargocentronpunctatissimum p=97-5,6.

Capros aper p=55-0,1. Antigonia capros p=97-3,4. Cyttus novaezelandiae p=9-3,4; 22-1,2; 95-1,2. Cyttus australis p=9-2,3. Cyttus traversi p=95-3,4; 97-2,3. Pseudocyttus maculatus p=94-5,6; 97-2,3. Oreosoma atlanticum p=

58-2,3; 71-0,1; 72-3,5; 94-4,5. Neocyttus rhomboidalis p=95-3,4; 97-2,3; 100-3,4. AUocyttus verrucosus p=33-0,l;

58-1,2; 99-7,8. AUocyttus niger p=33-0,l; 53-1,2. Parazen pacificus p=97-7,8. Cyttopsis roseus p=97-3,4. Zenion

hololepis p=33-0,l; 97-2,3. Capromimus abbreviatus p=97-3,4. Cyttomimus stelgis p=71-l,2; 72-3,4; 95-3,4.

Macrurocyttus acanthopodus p=22-l,2. Xenolepidichthys dalgleishi p=72-2,3. Zeusfaber p=9-2,3; 58-2,3; 95-1,2.

Zenopsis conchifer p=6-0,l; 95-1,2; 98-4,5,6. Zenopsis nebulosus p=6-0,1; 98-4,5.

Taxon

Melamphaes suborbitalis

Anoplogaster cornuta

Hoplostethus mediterraneus

Centrobervx affinis

Morone americana

Plectrypops lima

Sargocentron punctatissimum

Capros aper

Antigonia capros

Parahollardia lineata

Cyttus novaezelandiae

Cyttus australis

Cyttus traversi

Pseudocyttus maculatus

Oreosoma atlanticum

Neocyttus rhomboidalis

AUocyttus verrucosus

AUocyttus niger

Parazen pacificus

Cyttopsis roseus

Stethopristes eos

Zenion hololepis

Capromimus abbreviatus

Cyttomimus stelgis

Macrurocyttus acanthopodus

Xenolepidichthys dalgleishi

Grammicolepis brachiusculus

Zeusfaber

Zenopsis conchifer

Zenopsis nebulosus

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21100

00002

00002

00002

00000

00000

00000

00000

00000

00001

00000

00000

00010

00011

00001

70000

01100

01100

10020

10020

10020

11112

67890

11001

01000

01000

01000

01000

01000

00001

01000

11000

11000

11110

11110

11110

11110

11110

11110

11110

11110

11110

11110

11110

11111

11110

10110

11120

11111

11111

11110

11110

11110

22222

12345

40000

02002

40001

40001

40000

42002

40002

32000

40000

40001

Ipll2

12112

12112

70112

12112

12112

12112

12112

20112

11112

10112

22112

12112

12112

Ipll2

00112

02112

11112

10112

10112

22223

67890

11000

10000

10000

10000

11000

10000

10000

10000

11000

11000

00001

00001

00001

00110

00000

00110

00110

00220

00220

00110

00210

00210

00210

00210

00000

00001

00007

00210

00210

00210

33333

12345

00001

00001

10001

00000

00301

00100

00100

00311

00001

00211

10000

10000

10000

10000

10000

10000

lOpOO

lOpOO

10100

10000

10000

lOpOO

10200

10200

11101

11001

11001

10000

10000

10000

33334

67890

00010

00010

00010

00110

20110

00010

00110

01111

21111

01001

01011

11011

01011

01011

01011

11011

11011

11011

01001

01011

00011

00111

00011

00011

00701

01011

01011

11011

21001

21001

Characters and states

Morphometric

44444

12345

0-100

0-700

1-300

0-000

0-000

0-000

0-000

00000

00010

00000

70711

10111

70711

10710

10710

10710

10710

10710

10102

10112

10712

11100

10110

10110

71710

11110

71710

10210

10210

10210

44445

67890

15010

05310

05110

15010

15010

15010

15010

14010

15010

13001

03101

03101

03100

17001

11101

11001

11001

11701

01000

01201

01301

01001

02001

03000

13301

01000

15000

00101

00001

00101

55555

12345

00410

00010

00000

00000

00000

00000

00000

OOOOp

00000

20001

11211

11211

11211

11211

11111

11111

11111

llpll

11111

11211

21211

11211

21211

21211

21511

12211

12211

21211

21211

21211

55556

67890

*

00-00

0--00

00-20

00-10

00-10

00-20

00-21

00-21

00-21

00111

00-21

00-21

00121

00111

03pll

03111

03pll

03211

01-21

00121

00121

03121

02121

02-21

02221

02111

02111

13p31

13231

13231

66666

12345

-0030

--no

-0021

-0031

-0031

-0021

00021

1010-

10121

1110-

11111

11111

11111

1111?

11110

11111

11111

11111

0110-

11111

11111

11111

11111

11111

11111

11110

1111?

1110-

1110-

1110-

66667

67890

-0-11

---03

00013

00012

00022

00010

00010

00024

00025

---31

0107?

0007?

0207?

100??

1007?

1007?

1007?

100??

-0-22

-1-7?

-0-11

10112

101??

10012

---22

10112

00012

1007?

100??

100??

mil

12345

01000

04001

16001

13000

ppOOO

00000

00000

25001

37001

10000

14111

14111

72111

??101

pplOl

14101

12101

72101

23100

11110

11110

13101

24101

pplOl

12101

lplOl

13101

13101

13101

14101

1111%

67890

-0000

00004

10000

-1000

-0001

-1000

-0000

11002

11001

-0003

10000

11000

11000

10000

11000

11000

11000

11000

-0000

-0000

-0000

01000

01000

11000

0000?

00000

00000

00111

00111

00111

88888

12345

20001

21012

20221

20001

10001

10001

10001

20011

10011

20003

20101

20101

20101

20001

20001

20001

20001

20001

01200

21202

21202

10000

20220

20200

20000

20012

20012

20102

21102

21102

88889

67890

01000

00001

00400

00000

00000

00000

00000

00000

00000

20000

01010

00010

00010

00000

00000

00000

00000

00000

00000

01101

01101

00002

00002

00002

0300?

02002

02002

11201

13301

13301

99

12

00

00

00

00

00

00

00

00

00

00

00

10

00

01

01

01

01

01

00

00

00

00

00

00

00

00

00

00

20

00

Meristic

1111

99999990000

34567890123

22804001210

14707000220

0250p203420

01505p04110

03509613p31

1230p614310

1250p613210

01122513232

0104p413132

00240220162

32p32412021

32232412021

82p3p522321

8p33pl72431

6p432383421

75p3p39p421

643322p2421

75432383421

4343p411221

3233p331101

32334231101

1223p222221

3243p322221

22p33412221

54130350152

61511242121

92711342221

35p31654111

55p31p53031

56131p53031
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

CHARACTER 9.—Number of infraorbitals (well-developed

elements exclusive of lachrymal, dermosphenotic, and variable

rudiments): none (0); three or four (1); five or six (2); seven or

eight (3); nine or 10 (4); 11 or 12 (5).

CHARACTER 10.—Depth of most infraorbitals: relatively

slender and tubular (0); deep, with large pores and bridges or

open lacunae between upper and lower edges (1); deep, with

serrate vertical supporting flanges (2); absent (-).

There is no reason to believe that this multistate character is

an ordered transformation series; therefore, it was treated as

unordered throughout the analyses.

CHARACTER 11.—Dermosphenotic: an ossification distinctly

separate from sphenotic, sometimes relatively free from skull

(0); fused or highly integrated with sphenotic (1); absent as an

identifiable part of sphenotic (2).

CHARACTER 12.—Mouth size: large, alveolar process of premaxilla longer than lateral ethmoid depth (0); small, alveolar

process much less than lateral ethmoid depth (1).

CHARACTER 13.—Postmaxillary process: present (0); absent

C DCHARACTER 14.—Alveolar process of the premaxilla: simple

(0); ventrally indented to form pair of blunt lobes (1); deeply

bifurcated ventrally (2).

CHARACTER 15.—Ascending process of the premaxilla:

reaching to a level in front of orbit or about to front of orbit and

lateral ethmoid, no more than one-fifth into orbit (0); reaching

distinctly behind lateral ethmoid to about one-third into orbit

(1); reaching to about one-half into orbit (2); reaching to rear of

orbit (3).

CHARACTER 16.—Palatine teeth: present (0); absent (1).

CHARACTER 17.—Ectopterygoid teeth: present (0); absent

(1).

CHARACTER 18.—Articulation of palatine with cranium:

main axis of palatine relatively parallel, or only moderately oblique, to body axis and with fixed, dual articulation with lateral

ethmoid and ethmo-vomerine region (0); palatine usually oriented distinctly obliquely to axis of body and with a single,

pivotal, articulation with lateral ethmoid, resulting in considerable mobility (1).

CHARACTER 19.—Metapterygoid size and articulation: relatively large, length about three-quarters or more that of the

quadrate and articulating with it (0); reduced, length about onehalf or less of that of the quadrate and not articulating with it

(1); absent (2).

CHARACTER 20.—Symplectic ventral flange: absent (0);

present (1).

CHARACTER 21.—Dentary cartilages (on lateral surface of

dentary): absent or unconsolidated (0); two cartilages of moderate length, each attached anteriorly to dentary and lying sequentially one behind the other, first shorter than second (1);

two cartilages of moderate length, each attached anteriorly to

dentary and lying sequentially one behind the other, about

equal in size or first cartilage only slightly shorter than second

(2); well-developed, long complex consisting of two pieces,

posterior end of anterior piece connecting to anterior end of

second piece (3); single rod-like cartilage lying along lateral

surface of dentary' and attaching to it anteriorly (4). Treated as

unordered throughout analyses.

CHARACTER 22.—Serrations on lower border of dentary:

none (0); single barb near symphysis (1); multiple serrations behind symphysis (2). Treated as unordered throughout analyses.

CHARACTER 23.—Open gill slit between fourth and fifth ceratobranchials: present (0); absent (1).

CHARACTER 24.—Number of gills: 4 complete gills (one

complete gill of 2 hemibranchs on each complete gill arch), or

8 hemibranchs (0); 3Vi gills, or 7 hemibranchs, with no hemibranch on the rear of the fourth ceratobranchial (1). See following character (25) for comments on correlated features.

CHARACTER 25.—Number of series of gill rakers on branchial arches: 4'/2, a series present along rear of fourth gill slit (at

least dorsally), i.e., along anterior border of fifth ceratohyal (0);

4, no series along rear border of fourth gill slit (1); 3'/2, no series along posterior border of fourth ceratobranchial and none

along rear border of fourth gill slit (2).

We recognize that characters 23-25 may be correlated and

actually represent only a single character; however, because we

lack clear evidence supporting that assumption, we treat them

separately.

CHARACTER 26.—Uncinate process on first epibranchial: absent (0); present (1).

CHARACTER 27.—Interarcual cartilage: absent (0); present

(1).

CHARACTER 28.—Second pharyngobranchial suspensory

shaft: short or absent (0); moderately long (1); long, about onehalf to two-thirds length of first pharyngobranchial (2).

CHARACTER 29.—Third pharyngobranchial suspensory

shaft: short or absent (0); moderately long (1); long, almost as

long as shaft of second pharyngobranchial (2).

CHARACTER 30.—Position of upper surface of first basibranchial: level with dorsal surfaces of basihyal and second basibranchial, at least posteriorly (0); entirely below level of dorsal

surfaces of basihyal and second basibranchial (1).

CHARACTER 31.—Fourth upper pharyngeal toothplate:

present (0); absent (1).

CHARACTER 32.—Fifth ceratobranchial: toothed (0); toothless (1).

CHARACTER 33.—Beryciform foramen: present as completely enclosed opening (0); represented by deep groove along

lateral surface of ceratohyal, often onto dorsal hypohyal (1);

represented by deep concavity on dorsal surface of ceratohyal

(2); absent, no lateral groove and no deep dorsal concavity (no

deeper than ventral concavity) (3). Treated as unordered

throughout analyses.

CHARACTER 34.—Placement of heads of rear group of branchiostegal rays: over surface or along ventral edges of both the

ceratohyal and epihyal (0); clustered along ossified posterior

border of ceratohyal (1).
NUMBER 618 19

The distinction is somewhat subjective, as mentioned in the

"Previous Phylogenetic Analyses of Zeiforms" under Rosen

(1984).

CHARACTER 35.—Prominent notches on lower border of ceratohyal: present for heads of some branchiostegal rays in anterior group (0); absent (1).

CHARACTER 36.—Ceratohyal-epihyal articulation: exclusively through cartilage (0); through cartilage but with bony interdigitated articulations in some specimens, especially with increasing specimen size (1); bony interdigitated articulations in

all specimens at all sizes (2).

CHARACTER 37.—Depth of anterior end of epihyal: equal, or

about equal, to depth of adjacent part of ceratohyal (0); distinctly less deep than adjacent part of ceratohyal (1).

The distinction is somewhat subjective, as mentioned in the

"Previous Phylogenetic Analyses of Zeiforms" under Rosen

(1984).

CHARACTER 38.—Urohyal size: small to moderate, no

longer than ceratohyal (0); large, longer than ceratohyal (1).

Vertebral Column and Median Fins

CHARACTER 39.—First vertebra in caudal peduncle with a

modified neural or haemal spine: PU2 (0); PU3 (1).

CHARACTER 40.—Association of neural arch and spine of

first vertebra with skull: neural arch and spine not closely applied to skull (0); neural arch and spine closely applied to skull,

primarily to exoccipitals (1).

Johnson and Patterson (1993) noted certain differences between the caproid and zeiform conditions of this character, but

these differences do not necessarily refute homology of the feature in these groups.

CHARACTER 41.—Articulation of second and next few anterior abdominal vertebrae with skull-first vertebra: not flexible

(0); flexible, vertebrae linked laterally and ventrally by ligamentous bands, appearing as ventral straps by transmitted light

in lateral view (1).

CHARACTER 42.—Dorsal extension of neural spine of first

vertebra when neural arch and spine closely applied to skull:

neural spine extending only slightly, or not at all, dorsally

above its attachment to skull (0); neural spine with a long dorsal portion free from skull beyond curvature of supraoccipital

and exoccipitals (1); first neural arch and spine not closely applied to skull (-).

CHARACTER 43.—Proximal attachment of Baudelot's ligament: to basioccipital (0); to first vertebra (1); to exoccipitals

(2); to both basioccipital and first vertebra (3). Treated as unordered throughout analyses.

Our coding does not indicate polarity, and we accept Johnson

and Patterson's (1993) analysis showing that the primitive

acanthomorph condition is having the origin on the first vertebra (our state 1). See "Previous Phylogenetic Analyses of Zeiforms" under Johnson and Patterson (1993).

CHARACTER 44.—Neural spine orientation: all (or all but

first few) abdominal neural spines oriented posterodorsally (0);

several posterior abdominal and/or anterior caudal neural

spines oriented anterodorsally, or at least vertically (1).

CHARACTER 45.—Haemal arch and spine vacuities: no

prominent vacuities (0); vacuities of moderate size primarily in

haemal arches of many posterior abdominal vertebrae and often in those of anterior caudal vertebrae (1); large vacuities in

haemal arches or spines of many posterior abdominal vertebrae

and often in those of anterior caudal vertebrae (2). Treated as

unordered throughout the analyses.

CHARACTER 46.—Abdominal haemal spines: many vertebrae, especially posteriorly, with a transverse bony bridge below haemal canal and prominent process in the midline below

bridge (0); haemal arches with a transverse bony bridge below

haemal canal but no median spine below bridge, but short vertical projections may occur below bridge on each side (1).

CHARACTER 47.—Ossified ribs: present on most abdominal

vertebrae behind fourth (0); present only on last few abdominal

vertebrae (1); present only on a few middle abdominal vertebrae (2); absent (3); present on all abdominal vertebrae except

first (4); present on all abdominal vertebrae except first two (5).

Treated as unordered throughout analyses.

CHARACTER 48.—Ossified epineurals: present on most abdominal vertebrae or their ribs (0); present on a few anterior abdominal vertebrae (1); present on a few middle abdominal vertebrae (2); absent (3). Treated as unordered throughout analyses.

CHARACTER 49.—Length of the neural spine of PU2: long

(0); absent to short (1).

CHARACTER 50.—Hypurapophysis: present (0); absent (1).

CHARACTER 51.—Number of epurals: three (0); two (1); one

(2).

CHARACTER 52.—Articulation of proximal end of parhypural to urostylar centrum: strongly embracing centrum (0);

slightly removed from and not embracing centrum (1); laterally

expanded as a specialized peg on each side of parhypural, with

pegs fitting into sockets on each side of centrum (2). Treated as

unordered throughout analyses.

CHARACTER 53.—Degree of hypural fusion ("+" indicates

fused together): 4-6 separate hypural elements (0); hypurals 1

+ 2 fused to centrum and hypurals 3 + 4 free from centrum (1);

hypurals 1+2 + 3+4 fused to centrum (2); hypurals 1 + 2 and 3

+ 4 separate and both plates free from centrum (3); hypurals 1

+ 2 free from centrum and hypurals 3 + 4 + 5 either free or fused

to centrum (4); all hypurals fused to centrum and hypural 5 not

free (5). Treated as unordered throughout analyses.

CHARACTER 54.—Uroneural: present (0); absent (1).

CHARACTER 55.—Stegural (sensu Rosen, 1984): present (0);

absent (1).

CHARACTER 56.—Extra-caudal ossicle (sensu Fujita, 1990;

an extra bone between HPU2 and HPU3) in haemal spine of

PU2: absent in all specimens (0); present in at least some

specimens (1).
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

CHARACTER 57.—Dorsal-fin-spine locking mechanism: absent (0); present between first and second dorsal-fin spines (1);

present between second and third dorsal-fin spines (2); present

between first, second, and third dorsal-fin spines (3); no dorsalfin spines (-). Treated as unordered throughout analyses.

CHARACTER 58.—Number of groups of vacant interneural

spaces (when two or more spaces vacant): one (0), treated as

not applicable; two (1); three (2); four (3); only one or no

spaces vacant (-).

CHARACTER 59.—Number of dorsal-fin pterygiophores anterior to neural spine of fourth abdominal vertebra: none (0); two

(1); three (2); four (3).

CHARACTER 60.—Placement of first pterygiophore of

spinous dorsal fin: behind first interneural space (behind second or subsequent neural spines) (0); in first interneural space

(between first and second neural spines, or preneural space if

first neural arch and spine not closely applied to skull) and often slanted forward (1).

CHARACTER 61.—Position of base of first dorsal-fin pterygiophore in first interneural/preneural space: middle to rear, not

contacting skull or neural arch and spine of first vertebra (0);

front, contacting skull and first vertebra between sides of neural arch and spine (1); first pterygiophore not in first interneural

space (-).

CHARACTER 62.—Distal radials of spinous dorsal fin: large,

ossified (0); reduced, absent, or cartilaginous (1); dorsal-fin

spines absent (-). See comments under chain-link dorsal-fin

spine articulation in Johnson and Patterson (1993).

CHARACTER 63.—Symmetry of soft dorsal- and anal-fin

pterygiophores: asymmetrical (0); symmetrical (1).

CHARACTER 64.—Number of supraneurals: none (0); one

(1); two (2); three (3).

CHARACTER 65.—Cartilage at distal end of supraneural:

present (0); absent (1); no supraneurals (-).

CHARACTER 66.—Anal-fin-spine locking mechanism: absent, when two or more spines are present (0); present between

first and second spines (1); one or no spines (-).

CHARACTER 67.—Articulation of first anal-fin spine with

pterygiophore: unfused (0); fused in some populations or at

larger specimen sizes (1); fused in all specimens (2); anal-fin

spines absent (-).

We do not have early developmental evidence about this apparent fusion, and we infer it on the basis of adult comparative

morphology.

CHARACTER 68.—Length of second anal-fin spine: moderate

to long, more than one-half length of first spine (0); short, less

than one-half length of first spine (1); second spine absent (-).

CHARACTER 69.—Number of anal-fin pterygiophores in prehaemal space (anterior to haemal spine of first caudal vertebra): three (0); two (1); one (2); none (3); indeterminate (?).

This count is difficult to determine in many specimens because of the shortness of the haemal spines of the first and second caudal vertebrae and their close association along the top

rear of the large first anal-fin pterygiophore.

CHARACTER 70.—Number of anal-fin pterygiophores in first

interhaemal space (between haemal spines of the first and second caudal vertebrae): none (0); one (1); two (2); three (3); four

(4); five (5); indeterminate (?) (see character 69).

CHARACTER 71.—Number of anal-fin pterygiophores in second interhaemal space (between haemal spines of the second

and third caudal vertebrae): none (0); two (1); three (2); four

(3); indeterminate (?) (see character 69).

CHARACTER 72.—Number of anal-fin pterygiophores anterior to haemal spine of third caudal vertebra: three (0); four (1);

five (2); six (3); seven (4); eight (5); nine (6); 10 (7); 11 (8); indeterminate (?).

CHARACTER 73.—Dorsal-, anal-, and pectoral-fin rays:

branched (0); unbranched (1).

Paired Fin Girdles

CHARACTER 74.—Lateral flange on lowermost pectoral-fin

radial: absent (0); present (1).

CHARACTER 75.—Number of separate bony elements in

postcleithrum: two (0); one (1).

CHARACTER 76.—Flanges on the single postcleithrum: absent (0); one along posterior border, may be laterally flared,

(1); two postcleithra (-).

We have no evidence on whether the single postcleithrum results from the fusion of two postcleithra or from the loss of one

or the other postcleithrum.

CHARACTER 77.—Supracleithral serrations: absent (0);

present along posterior border, this border sometimes laterally

flared (1).

CHARACTER 78.—Ventral end of supracleithrum: simple (0);

deeply bifurcate (1).

CHARACTER 79.—Cleithral process above articulation with

postcleithrum: absent (0); present as prominent prong (1).

CHARACTER 80.—Extrascapulars: one long bone, sometimes

forming an open tube, more or less closely held to skull and integrated in line with the crest (often spiny) between the posttemporal and parietal (0); two tubular bones, not closely held to

skull except at larger specimen sizes (1); three more-or-less tubular bones (2); absent (3); one very small bone not integrated

into skull (4). Treated as unordered throughout analyses.

CHARACTER 81.— Pelvic fin position: about midway between anus and pectoral-fin base (0); slightly behind pectoralfin base (1); under or anterior to pectoral-fin base (2).

CHARACTER 82.—Pelvic-fin spines: present (0); absent (1).

CHARACTER 83.—Anterolateral processes of the medial

(lower) surfaces of the pelvic-fin rays: absent (0); present as

prongs from medial surfaces of ray bases (1); present as broad

flanges from ray bases (2). Treated as unordered throughout

analyses.

CHARACTER 84.—Pelvic-fin-ray serrations: absent (0);

present on crests on anterior or upper and/or lower posterior

surfaces of several rays (1); present on broad flanges from me-
NUMBER 618 21

dial (lower) surface of several rays (2). Treated as unordered

throughout analyses.

CHARACTER 85.—Basipterygia articulation: medial processes of basipterygia broadly overlap at level of pelvic fin (0);

in contact in midline of middle region but with little or no overlap (1): not in close contact in middle region but often in contact at anterior ends (2); tightly adherent or partially fused

along broad area of midline contact (3). Treated as unordered

throughout analyses.

CHARACTER 86.—Posterior process of the pelvis, behind pelvic-fin base: short to moderately long, shaped like a moderately

broad to broad plate or flattened shaft, usually slightly to distinctly obliquely oriented, with or without flanges and retrorse

projections (0); long and rod-like, moderately separated from

its opposite member along midline (1); long, tapering shaft in

contact or fused with its opposite member along midline (2).

Scales

CHARACTER 87.—Scales on most of body: moderate-sized to

small, spiny "ctenoid" (spinoid) scales (0); moderate-sized to

small, cycloid scales (1); scales greatly elongate vertically (2);

scales absent (excluding enlarged buckler-like scales), or with

only lateral-line scales (3). Treated as unordered throughout

analyses. For scale morphology in zeiforms and related taxa,

see Roberts (1993).

CHARACTER 88.—Buckler-like scales (see "Terminology"):

absent (0); present only from isthmus to anus (1); present midabdominally and from rear end of spinous dorsal fin to end of

soft dorsal-fin base (2); present mid-abdominally and from below spinous dorsal-fin base (usually from front to middle region) to end of soft dorsal-fin base (3); present mid-abdominally from pelvic fin to the anus (4). Treated as unordered

throughout analyses.

CHARACTER 89.—Scute-like scales (see "Terminology"): absent (0); present from isthmus to pelvic-fin base and sometimes

more posteriorly (1).

CHARACTER 90.—Scales along bases of dorsal- and anal-fin

rays: present, usually as a low sheath of scales lacking spiny

processes (0); absent, nearby scales without spiny projections

and not extending beyond lateral expansions of distal ends of

dorsal- and anal-fin pterygiophores (1); absent, but spiny processes present on scales alongside lateral expansions of distal

ends of dorsal- and anal-fin pterygiophores (2). Treated as unordered throughout analyses.

Miscellaneous

CHARACTER 91.—Hyperostosis: absent (0); present in supraoccipital and first dorsal-fin pterygiophore of some specimens (1); present in prepelvic scale bucklers (2). Treated as unordered throughout analyses.

CHARACTER 92.—Extended prejuvenile stage, with late ossification, thickened dermal connective tissue, expanded abdomen, and hillocks or cone-like scales: absent (0); present, (1).

Meristic Data

CHARACTER 93.—Total number of vertebrae: 26 or fewer

(0); 27 or 28 (1); 29 or 30 (2); 31 or 32 (3); 33 or 34 (4); 35 or

36 (5); 37 or 38 (6); 39 or 40 (7); 41 or 42 (8); 43 or more (9).

CHARACTER 94.—Number of abdominal vertebrae: nine or

fewer (0); 10(1); 11(2); 12(3); 13(4); 14(5); 15(6).

CHARACTER 95.—Number of vertebrae in caudal peduncle:

three (0); four (1); five (2); six (3); seven (4); eight (5); nine

(6); 10(7); 11 or more (8).

CHARACTER 96.—Number of principal caudal-fin rays: 16 or

more (0); 15 (1); 14 (2); 13 (3); 12 (4).

CHARACTER 97.—Number of procurrent caudal-fin rays (including number in both the dorsal and ventral sides, if different): none (0); one (1); two (2); three (3); four (4); five (5); six

(6); seven (7); eight (8); nine or more (9). See "Zeiform

Procurrent Caudal-Fin Rays" for discussion of specimen size

and development of procurrent rays.

CHARACTER 98.—Number of dorsal-fin spines: four or

fewer (0); five (1); six (2); seven (3); eight (4); nine (5); 10 or

more (6).

CHARACTER 99.—Total number of vacant interneural spaces

below dorsal-fin base, posterior to first dorsal-fin pterygiophore: none (0); one (1); two (2); three (3); four (4); five (5);

six (6); seven (7); eight (8); nine (9).

CHARACTER 100.—Number of anal-fin spines: none (0); one

(1); two (2); three (3); four (4); five (5).

CHARACTER 101.—Number of pectoral-fin rays: 11 or 12

(0); 13 or 14 (1); 15 or 16 (2); 17 or 18 (3); 19 or 20 (4); 21 or

22 (5).

CHARACTER 102.—Total number of pelvic-fin elements:

nine (0); eight (1); seven (2); six (3); five (4), not applicable to

listed taxa; four (5); three (6).

CHARACTER 103.—Number of branchiostegal rays: eight,

4 + 4 (0); seven, 3+4(1); six, 2 + 4 (2).

Phylogenetic Results

The following character optimizations are based solely on the

character distributions on the unordered consensus tree derived

from the reduced data set (Figure 4). Character-state changes

are given for the relevant nodes. When the character is further

derived or reversed in the members of the clade, this is noted.

When the character is further derived or reversed in members of

another clade convergently, the reader is directed to the relevant

optimization number for further details. Character states with

further change are only explicitly given where ambiguity is

possible (e.g., they are not given for binary characters).

Characters (e.g., optimization numbers 10-17) stated herein

to be convergent between zeiforms and the representative tetra-
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

odontiform (Parahollardia) and the two genera of caproids

{Capros and Antigonia) are potential synapomorphies of the

putative higher clades involving those taxa (as in Figures 1-3).

although these clades are not supported by the unordered analysis of the reduced data set (Figure 4).

MONOPHYLY OF ZEIFORMES (branch L in Figure 4 and the

Appendix).—The following nine consistent synapomorphies

diagnose a monophyletic zeiform clade.

1. Palatine usually oriented obliquely to axis of body and

with a single, pivotal articulation with lateral ethmoid,

resulting in considerable mobility (18-1); see also

Johnson and Patterson (1993).

2. Three and one-half gills, seven hemibranchs (24-1).

3. Dorsal-, anal-, and pectoral-fin rays unbranched (73-1);

convergent in one family (Monacanthidae) of tetraodontiforms not included in this study.

4. No uncinate process on first epibranchial (26-0).

5. Continuous median cartilage present below frontals, extending between ethmoid cartilage and pterosphenoids.

This is not included in the data set because the character could not be determined in many of our specimens of

zeiforms, but we agree with Johnson and Patterson

(1993) that this is a highly probable synapomorphy of zeiforms. For our documentation of this, see "Previous

Phylogenetic Analyses of Zeiforms" under Johnson and

Patterson (1993).

6. No open gill-slit between fourth and fifth branchial

arches (23-1).

7. Metapterygoid reduced in size and not articulating with

quadrate (19-1); see also Johnson and Patterson (1993);

metapterygoid lost in Macrurocyttus.

8. Parhypural slightly removed from and not embracing

urostylar centrum (52-1); see also Johnson and Patterson

(1993); further derived, to state 2, in Xenolepidichthys +

Grammicolepis.

9. Hypurals 1+2 + 3 + 4 fused and fused to centrum (53-2);

autapomorphy, to state 5, in Macrurocyttus; further derived, to state 1, in Oreosoma+ (Neocyttus + Allocyttus)

and Parazen.

The following 17 characters indicate zeiform monophyly but

exhibit homoplasy.

10. Long neural spine on PU2 (49-0); see also Rosen (1984);

convergent in Parahollardia.

11. Stegural absent (55-1); see also Rosen (1984); convergent in Parahollardia.

12. Distal radials of spinous dorsal fin absent or present only

as tiny cartilaginous or partially ossified structures under

the last few spines (62-1); see also Johnson and Patterson

(1993); convergent in Parahollardia.

Johnson and Patterson's (1993) analysis did not include tetraodontiforms, but we have found that both zeiforms and tetraodontiforms have this condition.

13. Hypurapophysis absent (50-1); convergent in Parahollardia; reversal in Cyttus traversi, Parazen, Cyttomimus,

and Xenolepidichthys + Grammicolepis.

14. Palatine teeth absent (16-1); convergent in Melamphaes,

Antigonia, and Parahollardia.

15. Neural spine closely applied to skull (40-1); convergent

in Capros, Antigonia, and Parahollardia.

16. Anterior end of epihyal distinctly less deep than adjacent

part of ceratohyal (37-1); convergent in Capros, Antigo

nia, and Parahollardia; reversal in Stethopristes, Zcnion + (Cyttomimus + Capromimus), and Macrurocyttus.

17. First pterygiophore of spinous dorsal fin inserted in first

interneural space, i.e., between first and second neural

spines (60-1); convergent in Sargocentron, Capros, Antigonia, and Parahollardia.

18. Three and one-half series of gill-rakers (25-2); convergent in Anoplogaster and Plectrypops + Sargocentron.

19. Prominent notches on lower border of ceratohyal (35-0);

convergent in Centroheryx and Plectrypops + Sargocentron; reversal in Macrurocyttus + {Xenolepidichthys +

Grammicolepis).

20. Uroneural absent (54-1); convergent in Melamphaes and

Anoplogaster.

21. Fourth upper pharyngeal toothplate absent (31-1); see

also Johnson and Patterson (1993); convergent in Hoplostethus.

22. Articulation of first few vertebrae with skull flexible,

vertebrae linked laterally and ventrally by ligamentous

bands (41-1); see also Johnson and Patterson (1993);

convergent in Hoplostethus.

23. Two dentary cartilages of moderate length, first shorter

than second (21-1); further derived, to state 2, in Parazen

and Zenion; further derived, to state 0, in Xenolepidichthys + Grammicolepis, and convergent in Anoplogaster.

24. Several posterior abdominal and/or anterior caudal neural spines oriented anterodorsally or at least vertically

(44-1); convergent in Antigonia; reversal in Parazen and

Zenion.

25. One supraneural (64-1); convergent in Anoplogaster;

further derived, to state 0, in Parazen and Zeus + Zenopsis, and convergent in Capros and Parahollardia.

26. Two epurals (51-1); reversal, to state 2, in Stethopristes

and {Zenion + (Capromimus + Cyttomimus)) + ((Macrurocyttus + (Xenolepidichthys + Grammicolepis)) + (Zeus

+ Zenopsis)), and convergent in Parahollardia; convergent secondary gain, to state 1, in Zenion and Xenolepidichthys + Grammicolepis.

The following three characters are optimized as supporting

the clade; however, their support is questionable because of unknown data or optimization ambiguity.

27. Baudelot's ligament attached to first vertebra (43-1); see

also Johnson and Patterson (1993); convergent in Melamphaes; further derived, to state 2, in Zeus + Zenopsis.
NUMBER 618 23

Because of the large number of taxa coded as unknown

(?), character 43-1 is questionable. For our documentation of this, see "Previous Phylogenetic Analyses of Zeifonns" under Johnson and Patterson (1993).

28. Vomerine teeth present (3-0); convergent in Centroberyx,

Morone, Plectrypops + Sargocentron, and Capros; reversal in Pseudocyttus, Oreosoma, and Macrurocyttus +

(Grammicolepis + Xenolepidichthys). Ambiguous optimization.

29. Infraorbitals relatively slender and tubular (10-0); convergent in Morone, Sargocentron, and Antigonia; further

derived, to state 2, in Zenion, and convergent in Anoplogaster and Hoplostethus; reversal, to state 1, in Oreosomatidae. Ambiguous optimization.

When optimized onto the tree, the following six characters

support this clade, but they are not used in the construction of

the tree, and this support is not used in the calculation of the

branch-support value.

30. Thirteen principle caudal-fin rays (96-3); further derived,

to state 1, in Xenolepidichthys + Grammicolepis.

31. Eight dorsal-fin spines (98-4); convergent in Antigonia;

further derived, to state 3, in all zeiforms except genus

Cyttus; further derived, to state 5, in Cyttus traversi; further derived, to state 1, in Pseudocyttus; further derived,

to state 2, in Allocyttus verrucosus, Stethopristes, Zenion,

and Xenolepidichthys, and convergent in Hoplostethus

and Parahollardia; reversal, to state 6, in Zeus + Zenopsis; reversal, to state 4, in Parazen and Cyttomimus.

32. Seven pelvic-fin elements (102-2); convergent in Anoplogaster and Hoplostethus; further derived, to state 0, in

Cyttopsis + Stethopristes; further derived, to state 5, in

Macrurocyttus; further derived, to state 1, in Zeus, and

convergent in Melamphaes, Centroberyx, and Plectrypops + Sargocentron; reversal, to state 3, in Pseudocyttus and Zeus + Zenopsis.

33. Seven branchiostegal rays (103-1); convergent in Morone; further derived, to state 2, in Macrurocyttus, and

convergent in Capros, Antigonia, and Parahollardia.

34. Two anal-fin spines (100-2); further derived, to state 1,

in Parazen + (Cyttopsis + Stethopristes), and convergent

in Melamphaes; further derived, to state 0, in Macrurocyttus, and convergent in Anoplogaster and Parahollardia; further derived, to state 4, in Zeus + Zenopsis, and

convergent in Centroberyx and Plectrypops; reversal, to

state 3, in Oreosoma, Neocyttus, Allocyttus niger, and

Zenopsis.

35. Thirty-one or 32 vertebrae (93-3); further derived, to

state 8, in Cyttus traversi and Oreosomatidae; further derived, to state 7, in Oreosoma + Neocyttus + Allocyttus;

further derived, to state 6, in Oreosoma, Allocyttus verrucosus, and Xenolepidichthys; further derived, to state 4,

in Parazen; further derived, to state 2, in Cyttomimus,

and convergent in Melamphaes; further derived, to state

5, in (Macrurocyttus + (Xenolepidichthys + Grammicolepis)) + (Zeus + Zenopsis); further derived, to state 9, in

Xenolepidichthys + Grammicolepis; further derived, to

state 1, in Zenion, and convergent in Anoplogaster and

Plectrypops + Sargocentron; reversal, to state 3, in Zeus.

Thus, zeiform monophyly is evidenced by 35 synapomorphies, of which 32 are informative (three other characters,

27-29, may support the clade). The Bremer value of 10, the

second highest in the analysis, indicates very strong support for

the monophyly of the Zeiformes.

MONOPHYLY OF Cyttus: CYTTOIDEI, CYTTIDAE (branch M in

Figure 4 and the Appendix).—Three consistent synapomorphies support the monophyly of the three species of Cyttus and

its higher categories.

36. Vacuities of moderate size in haemal arches of posterior

abdominal vertebrae and often in those of more anterior

caudal vertebrae (45-1).

37. Ascending process of premaxilla reaching back about

one-half into orbit (15-2).

38. Scute-like scales present from isthmus to pelvic-fin base

(89-1).

The following five characters exhibit homoplasy but nevertheless support the monophyly of Cyttus.

39. Upper surface of first basibranchial entirely below level

of dorsal surface of basihyal and second basibranchial

(30-1); convergent in Xenolepidichthys + Grammicolepis; see optimization number 140 for details.

40. Anterolateral processes of pelvic-fin rays present as

prongs from medial surfaces of ray bases (83-1); convergent in Zeus + Zenopsis.

41. Lateral flange present on lowermost pectoral-fin radial

(74-1); convergent in Cyttopsis + Stethopristes.

42. Ossified epineurals on a few anterior vertebrae (48-1);

convergent in Hoplostethus, Oreosoma, and Zeus + Zenopsis; see optimization number 158 for details.

43. No ossified ribs (47-3); convergent in Parahollardia.

Cyttomimus, and Macrurocyttus.

The following three characters may support a monophyletic

Cyttus and its higher categories, but some information is polymorphic or optimization is uncertain.

44. Multiple serrations on dentary behind symphysis (22-2);

convergent in Anoplogaster, Plectrypops, Capros, Oreosoma + Neocyttus + Allocyttus, Zenion + (Capromimus +

Cyttomimus), and Grammicolepis; further derived, to

state 1, in Cyttus novaezelandiae, and convergent in Cyttopsis and Zeus. Cyttus novaezelandiae assigned state 1

by mutation coding. Ambiguous optimization.

45. Many abdominal haemal spines with a prominent process

in midline below bridge under haemal canal (46-0); convergent in Anoplogaster, Hoplostethus, and (Parazen +

(Cyttopsis + Stethopristes)) + ({Zenion + (Capromimus +

Cyttomimus)) + ((Macrurocyttus + (Xenolepidichthys +

Grammicolepis)) + (Zeus + Zenopsis))); see optimization

number 77 for details. Ambiguous optimization.
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

46. Seven or 8 infraorbitals (9-3); polymorphic in Cyttus

australis (9-2,3) and C. novaezelandiae (9-3,4). Although specimens of both species exhibit state 3, other

specimens exhibit states 2 or 4; thus character may not

support clade.

When optimized onto the tree, the following two characters

support this clade, but they are not used in the construction of

the tree, and this support is not used in the calculation of the

branch-support value.

47. Eleven or 12 pectoral-fin rays (101-0); convergent in Zenopsis; further derived, to state 3, in Cyttus traversi, and

convergent in Plectrypops.

48. Five vertebrae in caudal peduncle (95-2); convergent in

Parahollardia, Zenion, and (Macrurocyttus + (Xenolepidichthys + Grammicolepis)) + (Zeus + Zenopsis); see

optimization number 127 for details.

The Bremer value of seven, one of the highest in the analysis, indicates very strong support for the monophyly of Cyttus

and its higher categories (Cyttoidei, Cyttidae).

MONOPHYLY OF Cyttus australis AND Cyttus traversi

(branch O in Figure 4 and the Appendix).—Within the genus

Cyttus, no consistent synapomorphies group Cyttus australis

with Cyttus traversi to the exclusion of Cyttus novaezelandiae.

One character, however, that has convergently arisen within

other zeiforms and within some of the non-zeiform taxa supports this group.

49. Serrations present along posterior border of supracleithrum (77-1); convergent in Centroberyx, Plectrypops,

Capros, Antigonia, Oreosoma + Neocyttus + Allocyttus,

and Zenion + (Cyttomimus + Capromimus).

This unnamed clade is supported by a single character,

which exhibits homoplasy. This is reflected in a Bremer value

of one, indicating only weak support for the monophyly of this

clade. Bray (1983) placed two of the three species of Cyttus,

C. australis and C. novaezelandiae, in the subgenus Cyttus and

the third, C. traversi, in the subgenus Rhombocyttus. Our evidence, although weak, is not in agreement with this subgeneric

grouping.

MONOPHYLY OF Pseudocyttus, Oreosoma, Neocyttus, Allocyttus, Parazen, Cyttopsis, Stethopristes, Zenion, Capromimus,

Cyttomimus, Macrurocyttus, Xenolepidichthys, Grammicolepis, Zeus, AND Zenopsis: ZEIOIDEI (branch R in Figure 4 and

the Appendix).—The remainder of the zeiforms form a monophyletic clade supported by seven characters. None of these

characters are consistent synapomorphies; they all exhibit a

certain amount of homoplasy, with several reversals or convergences.

50. Third pharyngobranchial suspensory shaft moderately

long (29-1); further derived, to state 2, in Allocyttus niger

and Parazen; reversal, to state 0, in Oreosoma and

Macrurocyttus + (Xenolepidichthys + Grammicolepis).

51. Anal-fin-spine locking mechanism between first and second spines (66-1); reversal in Grammicolepis; Parazen,

Cvttopsis, Stethopristes, and Macrurocyttus with 1 or no

spines (-) and cannot be coded for this character.

The following characters may support this clade, but optimization is uncertain.

52. Second pharyngobranchial suspensory shaft long (28-2);

further derived, to state 1, in Oreosomatidae and Cyttop

sis; reversal, to state 2, in Allocyttus niger; reversal, to

state 0, in Oreosoma and Macrurocyttus + (Xenolepidichthys + Grammicolepis). Ambiguous optimization.

53. Ossified ribs present only on last few abdominal vertebrae (47-1); further derived, to state 0, in Zeus + Zenopsis; further derived, to state 3, in Capromimus + Cyttomimus and Macrurocyttus, and convergent in Parahollardia

and Cyttus; further derived, to state 2, in Capromimus;

reversal, to state 5, in Grammicolepis. Ambiguous optimization.

When optimized onto the tree, the following characters support this clade, but they are not used in the construction of the

tree, and this support is not used in the calculation of the

branch-support value.

54. Seven dorsal-fin spines (98-3); further derived, to state 1,

in Pseudocyttus; further derived, to state 2, in Allocyttus

verrucosus, Stethopristes, Zenion, and Xenolepidichthys,

and convergent in Hoplostethus and Parahollardia; reversal, to state 6, in Zeus + Zenopsis; reversal, to state 4,

in Parazen and Cyttomimus.

55. Six vertebrae in caudal peduncle (95-3); convergent in

Plectrypops; further derived, to state 1, in Macrurocyttus

and Zenopsis nebulosus; further derived, to state 7, in

Macrurocyttus + (Grammicolepis + Xenolepidichthys),

and convergent in Anoplogaster; further derived, to state

4, in Oreosoma, Allocyttus niger, Parazen, and Capromimus; further derived, to state 2, in Zenion and (Macrurocyttus + (Xenolepidichthys + Grammicolepis)) +

(Zeus + Zenopsis); reversal, to state 5, in Xenolepidichthys. Occurrence of polymorphic state 1 in some specimens of Zeus and Zenopsis conchifer not relevant at this

level of analysis.

56. Three procurrent caudal-fin rays (97-3); convergent in

Cyttus traversi; further derived, to state 1, in (Macrurocyttus + (Xenolepidichthys + Grammicolepis)) +

(Zeus + Zenopsis); further derived, to state 0, in Macrurocyttus, which is convergent in Parahollardia; further derived, to state 4, in Stethopristes and Capromimus; reversal, to state 2, in Zenion and Oreosoma + Neocyttus +

Allocyttus; reversal, to state 3, in Neocyttus. Ambiguous

optimization.

The Bremer value of one indicates only weak support for the

monophyly of the Zeioidei.
NUMBER 618 25

MONOPHYLY OF Pseudocyttus, Oreosoma, Neocyttus, AND

Allocyttus: OREOSOMATIDAE (branch S in Figure 4 and the Appendix).—There is one consistent synapomorphy that supports

the monophyly of this family.

57. Extended prejuvenile stage present, with a unique combination of many derived morphologies (92-1).

There are also three characters that exhibit homoplasy but

nevertheless support the monophyly of the family.

58. Two dorsal-fin pterygiophores anterior to neural spine of

fourth abdominal vertebra (59-1); convergent in Centroberyx, Morone, Parahollardia, and Xenolepidichthys +

Gramm icolepis.

59. Most infraorbitals deep, with large pores and bridges or

open lacunae (10-1); reversal.

60. Second pharyngobranchial suspensory shaft moderately

long (28-1); convergent in Cyttopsis; reversal, to state 0,

in Oreosoma; convergent, to state 2, in Allocyttus niger.

When optimized onto the tree, the following four characters

support this clade, but they are not used in the construction of

the tree, and this support is not used in the calculation of the

branch-support value.

61. Nineteen or 20 pectoral-fin rays (101-4); convergent in

Hoplostethus.

62. Eight vacant intemeural spaces (99-8); further derived, to

state 7, in Pseudocyttus and Allocyttus verrucosus; autapomorphy, to state 9, in Neocyttus. Ambiguous optimization.

63. Forty-one or 42 vertebrae in total (93-8); convergent in

Cyttus traversi; further derived, to state 7, in Oreosoma +

Neocyttus + Allocyttus; further derived, to state 6, in

Oreosoma and A llocyttus verrucosus, and convergent in

Xenolepidichthys. Ambiguous optimization.

64. Fourteen abdominal vertebrae (94-5); convergent in Zeus

+ Zenopsis; further derived, to state 4, in Allocyttus verrucosus, and convergent in Anoplogaster and Macrurocyttus; see optimization number 163 for details. Ambiguous optimization.

The Bremer value of two indicates moderate support for the

monophyly of the family Oreosomatidae.

MONOPHYLY OF Oreosoma, Neocyttus, AND Allocyttus:

OREOSOMAT1NAE (branch U in Figure 4 and the Appendix).—

Seven characters, when optimized, support the monophyly of

this clade. None of these is a consistent synapomorphy, as all

have arisen convergently in other taxa.

65. Hypurals 1 +2 fused and fused to centrum and hypurals

3 + 4 fused and free from centrum (53-1); convergent in

Parazen.

66. Dorsal-fin-spine locking mechanism between first, second, and third spines (57-3); convergent in (Zenion +

(Capromimus + Cyttomimus)) + ((Macrurocyttus +

(Grammicolepis + Xenolepidichthys)) + (Zeus + Zenopsis))\ see optimization number 92 for details.

67. Dentary with multiple serrations behind symphysis (22-

2); convergent in Anoplogaster, Plectrypops, Capros,

Cyttus, Zenion + (Capromimus + Cyttomimus), and

Grammicolepis.

68. Serrations along posterior border of supracleithrum (77-

1); convergent in Centroberyx, Plectrypops, Capros, Antigonia, Cyttus australis + Cyttus traversi, and Zenion +

(Capromimus + Cyttomimus).

The following optimized character supports this clade because of the trichotomy involving Oreosoma, Neocyttus, and

Allocyttus.

69. Ceratohyal-epihyal articulation through cartilage but

with interdigitated articulation in some specimens (36-1);

convergent in Cyttus australis and Zeus. Character supports grouping of Neocyttus and Allocyttus. Ambiguous

optimization.

When optimized onto the tree, the following two characters

support this clade, but they are not used in the construction of

the tree, and this support is not used in the calculation of the

branch-support value.

70. Thirty-nine or 40 vertebrae (93-7); further derived, to

state 6, in Oreosoma and Allocyttus verrucosus, and convergent in Xenolepidichthys. Ambiguous optimization.

71. Two procurrent caudal-fin rays (97-2); reversal, with independent convergent reversal in Zenion; reversal, to

state 3, in Neocyttus, with independent convergent reversal in Cyttus traversi. Ambiguous optimization.

Thus, the support of one of these characters is not self-evident, but it is optimized as supporting the clade because of the

trichotomy. The Bremer value of two indicates moderate support for the monophyly of the subfamily Oreosomatinae.

MONOPHYLY OF Allocyttus verrucosus AND Allocyttus niger

(branch X in Figure 4 and the Appendix).—The monophyly of

this clade is supported by three homoplasious characters.

72. Five anal-fin pterygiophores anterior to haemal spine of

third caudal vertebra (72-2); convergent in Cyttus traversi and Macrurocyttus.

73. Beryciform foramen represented by deep grove along lateral surface of ceratohyal (33-1); convergent in Plectrypops + Sargocentron, Parazen, and Macrurocyttus.

Polymorphic in these two species of Allocyttus (33-0,1),

but character assigned state 1 by mutation coding.

74. Three groups of vacant intemeural spaces (58-2); convergent in (Macrurocyttus + (Xenolepidichthys + Grammicolepis)) + (Zeus + Zenopsis). Allocyttus verrucosus

assigned state 2 by mutation coding. Ambiguous optimization.

The Bremer value of one indicates only weak support for the

monophyly of this unnamed clade.

MONOPHYLY OF Parazen, Cyttopsis, Stethopristes, Zenion,

Capromimus, Cyttomimus, Macrurocyttus. Xenolepidichthys,
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Grammicolepis, Zeus, AND Zenopsis (branch A1 in Figure 4 and

the Appendix).—The monophyly of this clade is supported by

three homoplasious characters.

75. Six anal-fin pterygiophores anterior to haemal spine of

third caudal vertebra (72-3); convergent in Centroberyx;

further derived, to state 1, in Cyttopsis + Stethopristes,

and convergent in Melamphaes; further derived, to state

2, in Macrurocyttus, and convergent in Allocyttus and

Cyttus traversi; reversal, to state 4, in Capromimus and

Zenopsis nebulosus.

76. Basipterygia not in close contact in middle region (85-2);

convergent in Anoplogaster; further derived, to state 0, in

Parazen, Zenion + (Cyttomimus + Capromimus), and

Macrurocyttus.

The support from the next character for this clade is questionable because of ambiguity in optimization.

77. Many abdominal haemal spines with a prominent process

in midline below bridge under haemal canal (46-0); convergent in Anoplogaster, Hoplostethus, and Cyttus; reversal, to state 1, in Macrurocyttus + (Xenolepidichthys +

Grammicolepis); reversal, to state 0, in Xenolepidichthys.

Ambiguous optimization.

The Bremer value of two indicates moderate support for the

monophyly of this unnamed clade.

MONOPHYLY OF Parazen, Cyttopsis, AND Stethopristes:

PARAZENIDAE (branch B' in Figure 4 and the Appendix).—The

monophyly of this clade is supported by one consistent synapomorphy and four others that exhibit homoplasy.

78. Large haemal-spine vacuities in many posterior abdominal vertebrae (45-2).

79. Pelvic-fin spines absent (82-1); convergent in Anoplogaster and Zenopsis.

80. Two separate postcleithra (75-0); convergent in Melamphaes, Centroberyx, Morone, Plectrypops + Sargocentron. and Parahollardia.

81. Anterolateral processes of medial surfaces of pelvic-fin

rays present as broad flanges from ray bases (83-2); convergent in Hoplostethus and Capromimus + Cyttomimus.

When optimized onto the tree, the following character supports this clade, but it is not used in the construction of the tree,

and this support is not used in the calculation of the branchsupport value.

82. One anal-fin spine (100-1); convergent in Melamphaes.

The Bremer value of two indicates moderate support for the

monophyly of the family Parazenidae.

MONOPHYLY OF Cyttopsis AND Stethopristes: CYTTOPSINAE

(branch D' in Figure 4 and the Appendix).—One consistent synapomorphy supports the monophyly of this clade.

83. Buckler-like scales present from isthmus to anus (88-1).

The following four homoplasious characters support the

monophyly of this clade.

84. Four anal-fin pterygiophores anterior to haemal spine of

third caudal vertebra (72-1); convergent in Melamphaes.

85. Lateral flange on lowermost pectoral-fin radial (74-1);

convergent in Cyttus.

86. Scales on most of body cycloid and of moderate to small

size (87-1); convergent in Melamphaes, Cyttus novaezelandiae, and Zeus.

87. Scales absent along bases of dorsal- and anal-fin rays,

and nearby scales without spiny projections anu no, _.,

tending beyond lateral expansions of distal ends of pterygiophores (90-1); convergent in Anoplogaster and Zeus +

Zenopsis.

The following character may support the monophyly of this

clade, but its optimization is ambiguous.

88. No ossified epineurals (48-3); convergent in Anoplogaster and Macrurocyttus; autapomorphy, to state 2, in

Cyttopsis. Ambiguous optimization.

When optimized onto the tree, the following three characters

support this clade, but they are not used in the construction of

the tree, and this support is not used in the calculation of the

branch-support value.

89. Three vacant interneural spaces (99-3).

90. Nine pelvic-fin elements (102-0).

91. Thirteen or 14 pectoral-fin rays (101-1); convergent in

Centroberyx, Antigonia, Parahollardia, and {Macrurocyttus + (Grammicolepis + Xenolepidichthys)) + (Zeus +

Zenopsis); see optimization number 118 for details.

The Bremer value of three indicates good support for the

monophyly of the subfamily Cyttopsinae.

MONOPHYLY OF Zenion, Capromimus, Cyttomimus, Macrurocyttus, Xenolepidichthys, Grammicolepis, Zeus, AND Zenopsis (branch G' in Figure 4 and the Appendix).—All five of the

characters that support this clade exhibit homoplasy.

92. Dorsal-fin-spine locking mechanism between first, second, and third spines (57-3); convergent in Oreosoma*

Neocyttus + Allocyttus; further derived, to state 2, in

Capromimus + Cyttomimus and Macrurocyttus + (Grammicolepis + Xenolepidichthys).

93. Scales absent along bases of dorsal- and anal-fin rays,

but spiny processes present on scales alongside lateral

expansions of distal ends of pterygiophores (90-2); further derived, to state 1, in Zeus + Zenopsis, and convergent in Anoplogaster and Cyttopsis + Stethopristes.

The following two characters may support the monophyly of

the clade, but their optimization is ambiguous.

94. No flange present on the single postcleithrum (76-0);

convergent in Anoplogaster; reversal in Cyttomimus.

Ambiguous optimization.

95. One epural (51-2); convergent in Parahollardia and Stethopristes; reversal, to state 1, in Zenion and Xenolepidichthys + Grammicolepis. Ambiguous optimization.
NUMBER 618 27

When optimized onto the tree, the following character supports this clade, but it is not used in the construction of the tree,

and this support is not used in the calculation of the branchsupport value.

96. Five vacant interneural spaces (99-5); further derived, to

state 2, in Zenion + {Capromimus + Cyttomimus), and

convergent in Parahollardia and Cyttus traversi; further

derived, to state 4, in Xenolepidichthys + Grammicolepis;

reversal, to state 1, in Cyttomimus. Ambiguous optimization.

The Bremer value of one indicates only weak support for the

monophyly of this unnamed clade.

MONOPHYLY OF Zenion, Capromimus, AND Cyttomimus:

ZENIONTIDAE (branch H1 in Figure 4 and the Appendix).—All

eight of the characters that support this clade exhibit homoplasy.

97. Dentary with multiple serrations behind symphysis (22-

2); convergent in Anoplogaster, Plectrypops, Capros,

Cyttus, Oreosoma + Neocyttus + Allocyttus, and Grammicolepis.

98. Serrations present along posterior border of supracleithrum (77-1); convergent in Centroberyx, Plectrypops,

Capros, Antigonia, Cyttus australis + Cyttus traversi,

and Oreosoma + Neocyttus + Allocyttus.

99. Anterior end of epihyal equal, or about equal, to depth of

adjacent part of ceratohyal (37-0); reversal, with independent convergent reversals in Stethopristes and

Macrurocyttus.

The following four characters only questionably support this

clade.

100. Medial processes of basipterygia broadly overlap at level

of pelvic fin (85-0); convergent in Parazen and Macrurocyttus. Ambiguous optimization.

101. Beryciform foramen represented by deep concavity on

dorsal surface of ceratohyal (33-2); convergent in Parahollardia. Polymorphic (33) in Zenion for states 0 and 1

but not 2, thus distribution of states supports clade only

indirectly, through homoplasy. Ambiguous optimization.

102. Alveolar process of premaxilla ventrally indented (14-1);

reversal in Cyttomimus. Ambiguous optimization.

103. Second anal-fin spine short (68-1); convergent in Xenolepidichthys; reversal in Cyttomimus. Ambiguous optimization.

When optimized onto the tree, the following character supports this clade, but it is not used in the construction of the tree,

and this support is not used in the calculation of the branchsupport value.

104. Two vacant interneural spaces (99-2); convergent in

Parahollardia and Cyttus traversi; reversal, to state 1, in

Cyttomimus.

The Bremer value of two indicates moderate support for the

monophyly of the family Zeniontidae.

MONOPHYLY OF Capromimus AND Cyttomimus (branch J' in

Figure 4 and the Appendix).—All six characters that support

this clade exhibit homoplasy.

105. Ascending process of premaxilla reaching distinctly behind lateral ethmoid to about one-third into orbit (15-1);

convergent in Parazen.

106. Three anal-fin pterygiophores in second interhaemal

space (71-2); convergent in Morone, Capros, and Parazen.

107. Seven anal-fin pterygiophores anterior to haemal spine of

third caudal vertebra (72-4); reversal, with independent

convergent reversal in Zenopsis nebulosus.

108. Anterolateral process of medial surfaces of pelvic-fin

rays present as broad flanges (83-2); convergent in Hoplostethus and Parazen + {Capromimus + Cyttomimus).

The following two characters only questionably support this

clade.

109. No ossified ribs (47-3); convergent in Parahollardia,

Cyttus, and Macrurocyttus; autapomorphy, to state 2, in

Capromimus. Ambiguous optimization.

110. Dorsal-fin-spine locking mechanism between second and

third spines (57-2); convergent in Macrurocyttus +

{Grammicolepis + Xenolepidichthys). Ambiguous optimization.

The Bremer value of two indicates moderate support for the

monophyly of this unnamed clade.

MONOPHYLY OF Macrurocyttus, Xenolepidichthys, Grammicolepis, Zeus, AND Zenopsis (branch M' in Figure 4 and the Appendix).—All eight characters that support the monophyly of

this clade exhibit homoplasy.

111. Scales absent (excluding buckler-like scales), or with

only lateral-line scales (87-3); further derived, to state 2,

in Xenolepidichthys + Grammicolepis; further derived, to

state 1, in Zeus, and convergent in Melamphaes, Cyttus

novaezelandiae, and Cyttopsis + Stethopristes.

112. Basisphenoid absent (2-2); convergent in Melamphaes

and Parahollardia; further derived, to state 1, in

Xenolepidichthys + Grammicolepis.

113. Lachrymal moderately deep (8-1); convergent in

Plectrypops + Sargocentron; further derived, to state 2, in

Zenopsis.

The support from the next character is questionable.

114. Three groups of vacant interneural spaces (58-2); convergent in Allocyttus; reversal, to state 1, in Xenolepidichthys + Grammicolepis. Ambiguous optimization.

When optimized onto the tree, the following four characters

support this clade, but they are not used in the construction of

the tree, and this support is not used in the calculation of the

branch-support value.

115. One procurrent caudal-fin ray (97-1); further derived, to

state 0, in Macrurocyttus, and convergent in Parahollardia.
28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

116. Thirty-five or 36 vertebrae (93-5); further derived, to

state 9, in Xenolepidichthys + Grammicolepis; further derived, to state 6, in Xenolepidichthys, and convergent in

Oreosoma and Allocyttus verrucosus; reversal, to state 3,

in Zeus.

117. Five vertebrae in caudal peduncle (95-2); convergent in

Parahollardia, Cyttus, and Zenion; further derived, to

state 7, in Macrurocyttus + {Xenolepidichthys + Grammicolepis), and convergent in Anoplogaster; further derived, to state 1, in Macrurocyttus and Zenopsis nebulosus, and convergent in Capros; reversal, to state 5, in

Xenolepidichthys.

118. Thirteen or 14 pectoral-fin rays (101-1); convergent in

Centroberyx, Antigonia, Parahollardia, and Cyttopsis +

Stethopristes; further derived, to state 0, in Zenopsis, and

convergent in Cyttus; reversal, to state 2, in Grammicolepis.

The Bremer value of one indicates only weak support for the

monophyly of this unnamed clade.

MONOPHYLY OF Macrurocyttus, Xenolepidichthys, AND

Grammicolepis: GRAMMICOLEPIDIDAE (branch N1 in Figure 4

and the Appendix).—One consistent synapomorphy supports

the monophyly of this clade.

119. Fifth ceratobranchial toothless (32-1).

The following five homoplasious characters support the

monophyly of this clade.

120. Neural spine of first vertebra with long dorsal portion

free from skull beyond curvature of supraoccipital and

exoccipitals (42-1); convergent in Zenion.

121. Vomerine teeth absent (3-1); reversal, with independent

convergent reversals in Pseudocyttus and Oreosoma.

122. Second pharyngobranchial suspensory shaft short or absent (28-0); reversal, with independent convergent reversal in Oreosoma.

123. Third pharyngobranchial suspensory shaft short or absent

(29-0); reversal, with independent convergent reversal in

Oreosoma.

124. No prominent notches on lower border of ceratohyal

(35-1); reversal.

The following two characters are of questionable support.

125. Most haemal arches of abdominal vertebrae with transverse bony bridge below haemal canal but no median

spine below bridge (46-1); reversal, to state 0, in Xenolepidichthys. Ambiguous optimization.

126. Dorsal-fin-spine locking mechanism between second and

third dorsal-fin spines (57-2); convergent in Capromimus

+ Cyttomimus. Ambiguous optimization.

When optimized onto the tree, the following character supports this clade, but it is not used in the construction of the tree,

and this support is not used in the calculation of the branchsupport value.

127. Ten vertebrae in caudal peduncle (95-7); convergent in

Anoplogaster; further derived, to state 1, in Macrurocyttus, and convergent in Zenopsis nebulosus; reversal, to

state 5, in Xenolepidichthys.

The Bremer value of four indicates strong support for the

monophyly of the family Grammicolepididae.

MONOPHYLY OF Xenolepidichthys AND Grammicolepis:

GRAMMICOLEPIDINAE (branch P' in Figure 4 and the Appendix).—Three consistent synapomorphies support the monophyly of this clade.

128. Proximal end of parhypural laterally expanded a^ .,, wcialized pegs, with pegs fitting into sockets on sides of

centrum (52-2).

129. Scales on most of the body greatly elongate vertically

(87-2).

130. Basisphenoid present as a short shaft at front of roof of

posterior myodome (2-1).

The following eight homoplasious characters support the

monophyly of this clade.

131. Mouth small (12-1); convergent in Antigonia and Parahollardia.

132. Symplectic ventral flange present (20-1); convergent in

Melamphaes, Sargocentron, and Zenion.

133. Two dorsal-fin pterygiophores anterior to neural spine of

fourth abdominal vertebra (59-1); convergent in Centroberyx, Morone, Parahollardia, and Oreosomatidae.

134. Pelvic-fin-ray serrations on crests on anterior or upper

and/or lower posterior surfaces of several rays (84-1);

convergent in Anoplogaster, Capros, and Antigonia.

135. Postmaxillary process absent (13-1); reversal, and convergent in Centroberyx and Parahollardia.

136. Hypurapophysis present (50-0); reversal, and convergent

in Cyttus traversi, Parazen, and Cyttomimus.

137. Two epurals (51-1); reversal, and convergent in Zenion.

138. Dentary cartilage absent or unconsolidated (21-0); convergent in Anoplogaster.

The following two characters are optimized as supporting the

clade, but only questionably.

139. Two groups of vacant interneural spaces (58-1); reversal.

Ambiguous optimization.

140. First basibranchial entirely below level of dorsal surfaces

of basihyal and second basibranchial (30-1); convergent

in Cyttus; Grammicolepis unknown. Ambiguous optimization.

When optimized onto the tree, the following three characters

support this clade, but they are not used in the construction of

the tree, and this support is not used in the calculation of the

branch-support value.

141. Fifteen caudal-fin rays (96-1).

142. Four vacant interneural spaces (99-4).

143. Forty-three or 44 vertebrae (93-9); further derived, to

state 6, in Xenolepidichthys, and convergent in Oreosoma

and Allocyttus verrucosus. Ambiguous optimization.

Even with the uncertainty of support from several of the

above characters, the Bremer value of eight is the third highest
NUMBER 618 29

in the entire cladogram, indicating very strong support for the

monophyly of the subfamily Grammicolepidinae.

MONOPHYLY OF Zeus AND Zenopsis: ZEIDAE (branch S1 in

Figure 4 and the Appendix).—The following nine consistent

synapomorphies support the monophyly of this family.

144. Otolith tiny, trilobed (7-1).

145. Alveolar process of premaxilla deeply bifurcate ventrally

(14-2).

146. Baudelot's ligament attached to exoccipitals (43-2).

147. Extra-caudal ossicle in haemal spine of PU2 of at least

some specimens (56-1).

148. Four dorsal-fin pterygiophores anterior to neural spine of

fourth abdominal vertebra (59-3).

149. Ventral end of supracleithrum deeply bifurcate (78-1).

150. Ossified ribs on most abdominal vertebrae behind fourth

vertebra (47-0).

151. Cleithral processes a prong above articulation with postcleithrum(79-l).

152. Posterior process of pelvis long, rod-like, moderately separated from its opposite member along midline (86-1).

The following five homoplasious characters support the

monophyly of this clade.

153. Dermosphenotic fused or closely consolidated with sphenotic (11-1); convergent in Capros.

154. Two extrascapular bones, not held closely to skull (80-1);

convergent in Antigonia and Morone.

155. Anterolateral processes of pelvic-fin rays present as

prongs from medial surfaces of ray bases (83-1); convergent in Cyttus.

156. Scales absent along bases of dorsal- and anal-fin rays,

nearby scales without spiny projections (90-1); convergent in Anoplogaster and Cyttopsis + Stethopristes.

157. No supraneurals (64-0); convergent in Capros, Parahollardia, and Parazen.

The following five characters are optimized as supporting

the monophyly of the group, but only questionably.

158. Ossified epineurals present on a few anterior abdominal

vertebrae (48-1); convergent in Hoplostethus, Cyttus, and

Oreosoma; reversal, to state 0, in Zenopsis conchifer.

Ambiguous optimization.

159. Bony interdigitated articulations between ceratohyal-epihyal in specimens of all sizes (36-2); convergent in Morone and Antigonia; further derived, to state 1, in Zeus,

and convergent in Cyttus australis and Oreosoma +

Neocyttus + Allocyttus. Ambiguous optimization.

160. Buckler-like scales present mid-abdominally and from

below front to middle region of spinous dorsal-fin base to

end of soft dorsal-fin base (88-3); autapomorphy, to state

2, in Zeus. Ambiguous optimization.

161. Supraocular serrations on frontal absent (6-1); convergent in Melamphaes, Morone, Sargocentron, and Parahollardia; polymorphic in Zenopsis (states 0, 1).

162. Eleven or 12 infraorbitals (9-5); further derived, to state

4, in Zenopsis conchifer; polymorphic in Zeus (states 2,

3). Character does not clearly support clade without optimization.

When optimized onto the tree, the following four characters

support this clade, but they are not used in the construction of

the tree, and this support is not used in the calculation of the

branch-support value.

163. Fourteen abdominal vertebrae (94-5); convergent in

Oreosomatidae; autapomorphy, to state 6, in Zenopsis

nebulosus.

164. Four anal-fin spines (100-4); convergent in Centroberyx

and Plectrypops; reversal, to state 3, in Zenopsis, with independent convergent reversals in Oreosoma and Allocyttus niger. Ambiguous optimization.

165. Six elements in pelvic-fin (102-3); reversal, with independent convergent reversal in Pseudocyttus; further derived, to state 1, in Zeus, and convergent in Melamphaes,

Centroberyx, and Plectrypops + Sargocentron. Ambiguous optimization.

166. Ten or more dorsal-fin spines (98-6); reversal; polymorphic in Zenopsis (Z. nebulosus, states 4, 5; Z. conchifer

states 4, 5, 6).

The Bremer value of 12, which is the highest in the analysis,

indicates very strong support for the monophyly of the family

Zeidae.

MONOPHYLY OF Zenopsis conchifer AND Zenopsis nebulosus

(branch U1 in Figure 4 and the Appendix).—One consistent synapomorphy supports the monophyly of Zenopsis.

167. Lachrymal slender (8-2).

The following two homoplasious characters support the

monophyly of this clade.

168. First vertebra in caudal peduncle with modified haemal

or neural spine PU2 (39-0); convergent in Parahollardia,

Parazen, and Macrurocyttus.

169. Pelvic-fin spines absent (82-1); convergent in Anoplogaster and Parazen + (Cyttopsis + Stethopristes).

When optimized onto the tree, the following two characters

support this clade, but they are not used in the construction of

the tree, and this support is not used in the calculation of the

branch-support value.

170. Eleven or 12 pectoral-fin rays (101-0); convergent in

Cyttus.

171. Three anal-fin spines (100-3); reversal, with independent

convergent reversals in Oreosoma and Allocyttus niger.

The Bremer value of two indicates moderate support for the

monophyly of this unnamed clade.

Previous Phylogenetic Analyses ofZeiforms

A number of workers have contributed to the study of the

phytogeny of zeiforms. Roman numerals in brackets distinguish the characters used in those previous analyses of zeiform
30 SMITHSONIAN CONTRIBUTIONS TC ZOOLOGY

relationships from the characters and their states described

herein, which are listed as Arabic numerals in parentheses.

Rosen (1984:31) listed seven characters (not eight, as stated

on p. 44 of that work) as indicating monophyly of caproids, zeiforms, and tetraodontiforms. We include six of these characters in our study of the relationships of these groups. Our analysis based on the unordered reduced data set (Figure 4) neither

supports nor rebuts this clade, because of the large polytomy

among the outgroups, but the other three analyses (Figures

1-3) do support it, and we offer the following comments on

Rosen's (1984:31) characters, using the same numerical

scheme that he employed.

[I]. "Saccular otolith (sagitta) higher than long and constricted fore and aft in the region of the sulcus." This characterization of overall shape does not apply to many zeiforms or tetraodontiforms, and among caproids it applies to Capros but not

to Antigonia. Moreover, the significance of the gross shape of

otoliths is very difficult to interpret and can usually be employed only in conjunction with detailed study of surface features. Many zeiforms and tetraodontiforms have a relatively

rounded or oval-shaped sagitta, similar to that of beryciforms.

Otolith shape corresponds to our character (7), in which the obviously derived state of reduced size and bow-tie shape supports only the clade of Zeus + Zenopsis. Therefore, we reject

the otolith shape character of zeiforms in general as evidence

of monophyly at Rosen's ordinal level.

[II]. "Dorsal and anal fin radials symmetrical,..." This corresponds to our character (63), which would be a synapomorphy of this putative clade of caproids, zeiforms, and tetraodontiforms; however, it occurs convergently in Anoplogaster.

[III]. "Radial supporting first dorsal fin spine robust, columnar, inserted ventrally between the neural arches of the first

vertebrae [sic] and the occipital bones of the skull to which

they are firmly united." Because the first pterygiophore does

not lie in the space between the skull and the first neural arch in

all the taxa studied herein, it is more appropriate to separate

Rosen's character into two distinct characters, as we have done

in our characters (60) and (61). The placement of the first

pterygiophore of the spinous dorsal-fin (60) supports this putative clade; however, it is convergently derived in Sargocentron.

The position of the base of the first dorsal-fin pterygiophore in

the first interneural space (61) would also be a synapomorphy

of this putative clade of caproids, zeiforms, and tetraodontiforms, but there is a secondary loss of contact between the

pterygiophore and the neural arch and spine in Parazen.

[IV]. "Parasphenoid with a tunnel-like opening to the posterior myodome." This is not discussed in the text by Rosen. The

derived condition of this character (4) applies to Morone, caproids, and tetraodontiforms. As optimized on our cladogram

(Figure 4), the presence of a tunnel-like opening in this area

does not support the clade, and this derived condition is not

present in any of the zeiform taxa.

[V]. "Interoperculum elongate, bladelike or feather-shaped."

There is so much variation in the shape of the interopercle

among zeiforms that this characterization applies only to some

taxa. We do not think that caproids share the stated condition,

although Parahollardia does. We find no way to characterize

the variation in the shape of the interopercle in a way that

would be useful in defining any of the three groups, let alone as

a monophyletic assemblage.

[VI]. "Premaxillary ascending and articular processes elongate and intimately united above a distinct neck that separates

them from a very short alveolar process." The variety of premaxillary shapes in zeiforms is so great that we find no unequivocal way to characterize the configuration of the premaxilla by region. Rosen's own figures of premaxillary shape

(1984, figs. 27-29) in selected species of caproids, zeiforms,

and tetraodontiforms do not all conform to his definition.

Rosen and Patterson (1990, figs. 12-15) and Johnson (1982,

figs. 2-20) show premaxillae similar to those of zeiforms in

many groups of acanthomorphs. We were able, however, to

code the extent of the elongation of the ascending process of

the premaxilla (15) for all taxa. This character is not a synapomorphy of zeiforms or any other higher categories. Moreover,

we are unable to characterize premaxillary shape in any way

that suggests it is a synapomorphy of the putative clade of caproids, zeiforms, and tetraodontiforms.

[VII]. "Caudal fin with 15 or fewer principal rays." This is

our character (96). We agree that this character occurs, although not uniquely among acanthomorphs, in each of the caproids, zeiforms, and tetraodontiforms and is supportive of this

putative clade.

We accept four (II, VII, and the two characters within III) of

Rosen's characters as support for this putative clade of caproids, zeiforms, and tetraodontiforms, but only Rosen's character VII is a consistent synapomorphy of the group within the

species in our data set. To this we add that the neural arch and

spine of the first vertebra is closely applied to the skull (40).

Thus, two consistent synapomorphies support the putative

clade of caproids, zeiforms, and tetraodontiforms.

We emphasize that although the present study is unable to

draw any definitive conclusion regarding the status of this putative clade of caproids, zeiforms, and tetraodontiforms, there

are several characters that do support it (Figures 1-3).

Zehren (1991:61 A), in a brief one paragraph abstract, stated

that caproids are monophyletic based on "seven apomorphies,

including ornamentation of the preopercle, frontal and supracleithrum," without listing all seven characters (but with the

three ornamentation characters counted separately; S. Zehren,

pers. comm., 16 Oct 1998). We are unable to find any consistent synapomorphies that define this clade of two genera (Capros and Antigonia). Other derived features, exhibiting homoplasy, support the caproid clade in three (Figures 1-3) of our

four analyses; however, this clade is not supported in the analysis (Figure 4) that we propose as the most rational.

Rosen (1984:31-32) gave four putative synapomorphies (not

five, as stated on p. 44) of tetraodontiforms and zeiforms.
NUMBER 618 31

[VIII]. "Caudal skeleton with a full spine on the second

preural centrum and a greatly reduced first uroneural with no

stegural." We regard the first (49) and third (55) of these as

valid possible synapomorphies for the putative clade of zeiforms and tetraodontiforms, albeit hardly unique to these fishes.

A relatively well-developed uroneural (54) is present among

tetraodontiforms in triacanthoids and in the primitive tetraodontoid Triodon and thus would not support this putative clade.

[IX]. "Greatest depth of caudal peduncle less than 13 percent

of standard length." The statement itself is generally true for

both zeiforms and tetraodontiforms, but it is also true for some

members of the outgroups (e.g., Centroberyx 12%; Capros

11%-12%). Thus, we have little confidence in this character as

evidence for higher-level relationships among these taxa.

[X]. "Hyoid apparatus with the posterior ceratohyal [=our

epihyal] very short and shallower than the adjoining part of

the anterior ceratohyal; the bases of the four posterior branchiostegals clustered on the posteroventral margin of the ossified anterior ceratohyal." The first part of this character (37)

could be a synapomorphy of this putative clade, although the

distinction is somewhat subjective, and, because in our interpretation the condition is also present in Capros and Antigonia, it would be a synapomorphy at a higher level (but we

stress the subjectivity of the character states). We also find the

positioning of the branchiostegal bases to be somewhat subjective, and in our interpretation (34), Rosen's derived condition seems to characterize Capros and Parahollardia, but not

Antigonia and zeiforms.

[XI]. "Interoperculum long and very slender, acinaciform,

and only loosely bound to the preoperculum by connective tissue." We find this subcondition of Rosen's character V to be

just as variable within zeiforms and tetraodontiforms as the

other interopercular shape characterization, and we find no useful way to define it as a shared derived feature of the two

groups.

Rosen (1984:33) listed six synapomorphies of tetraodontiform taxa.

[XII]. "No anal spines." This condition (100) supports the

monophyly of tetraodontiforms, but this loss is convergent in

Anoplogaster and Macrurocyttus.

[XIII]. "Caudal fin with 12 or fewer principal rays." The

condition of 15 or fewer caudal-fin rays (96) supports a putative

clade of caproids, zeiforms, and tetraodontiforms. Within this

clade, tetraodontiforms can be characterized as having 12 or

fewer principal rays; however, this condition is convergent in

Antigonia.

[XIV]. "No infraorbitals." This condition (9) is a synapomorphy of tetraodontiforms, but the infraorbitals are absent

convergently in Macrurocyttus.

[XV]. "No parietals." As with the infraorbitals, the lack of

parietals (1) is a synapomorphy of tetraodontiforms, but the parietals are absent convergently in Macrurocyttus.

[XVI]. "Small, slitlike gill opening just anterior to the pectoral fin base." We accept this condition as a consistent synapomorphy of tetraodontiforms (no character number is assigned

to this feature because it is autapomorphic in our data set for

Parahollardia; see "Analytical Protocols").

[XVII]. "Pelvic girdles closely joined along their lengths and

the pelvic-fin rays reduced or absent." The joining of the pelvic

girdle along its length (85) is a synapomorphy of tetraodontiforms. We regard the reduction or loss of the pelvic-fin ray as a

separate character, but it was not used in the present study because it is autapomorphic in our data set for Parahollardia.

Thus, of the six synapomorphies listed by Rosen for tetraodontiforms, we find that only two are non-homoplasious synapomorphies, but they represent three synapomorphies because

Rosen's character XVII should be divided into two separate

characters.

Rosen provided a single putative synapomorphy to link the

ostraciid boxfishes with tetraodontoids. Klassen (1995) pointed

out that this single character is far outweighed by the 20 synapomorphies that ostracioids share with balistoids and by the 15

synapomorphies uniting tetraodontoids that are not found in ostracioids, citing the evidence presented in Winterbottom (1974,

not referred to by Rosen) and Winterbottom and Tyler (1983).

Rosen (1984) did not propose any synapomorphies for zeiform

taxa.

Regarding tetraodontiforms, we note that nearly all of the 32

character states identified in the present study as autapomorphies of the representative tetraodontiform, Parahollardia lineata, in branch K. of Figure 4 and the Appendix, likewise are

synapomorphies for the clade comprising all of the extant families of tetraodontiforms, or of the putative zeiform + tetraodontiform clade, or of that clade plus caproids. Many of these tetraodontiform apomorphies involve reductions, simplifications, or

loss of structures. These are as follows, with those that are potential synapomorphies of the putative larger clade involving

zeiforms, and in many cases caproids, marked with an asterisk

after the character state: parietal absent (1-1); basisphenoid absent (2-2); parasphenoid opening into posterior myodome (4-

1); supraocular serrations on frontal absent (6-1); infraorbitals

absent (9-0); dermosphenotic absent (11-2); mouth small (12-

1); postmaxillary process absent (13-1); palatine teeth absent

(16-1); four series of gill rakers (25-1); interarcual cartilage

present (27-1); beryciform foramen represented by deep concavity on dorsal surface of ceratohyal (33-2); heads of branchiostegal rays clustered along posterior border of ceratohyal

(34-1); anterior end of epihyal distinctly less deep than adjacent part of ceratohyal (37-1*); PU2 first caudal peduncle vertebra with modified haemal or neural spines (39-0); neural arch

and spine of first vertebra closely applied to skull (40-1 *); ossified ribs absent (47-3*); PU2 neural spine long (49-0*); hypurapophysis absent (50-1*); one epural (51-2); stegural absent

(55-1*); two dorsal-fin pterygiophores anterior to fourth neural

spine (59-1); first dorsal-fin pterygiophore inserted in first interneural (or preneural) space (60-1*); distal radials of spinous

dorsal fin absent (62-1*); supraneurals absent (64-0); no analfin pterygiophores in prehaemal space (69-3); one anal-fin
32

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

pterygiophore in first interhaemal space (70-1); three anal-fin

pterygiophores anterior to third caudal haemal spine (72-0);

two elements in postcleithrum (75-0); extrascapulars absent

(80-3); basipterygia tightly adherent or fused along broad area

of midline contact (85-3); and long, tapering shaft-like posterior process of pelvis (86-2). Two other characters not listed

under branch K are higher-level potential synapomorphies of

the putative caproid + zeiform + tetraodontiform clade, these

being base of first dorsal-fin pterygiophore in contact with the

skull and first vertebra (61-1*), and symmetrical radials (63-

1*). Additionally, the following meristic features are apomorphies of Parahollardia and represent synapomorphies for the

clade comprising all of the extant families of tetraodontiforms:

nine or fewer (actually eight in Parahollardia) abdominal vertebrae (94-0); 12 caudal-fin principal rays (96-4); no procurrent

caudal-fin rays (97-0); six dorsal-fin spines (98-2); two vacant

interneural spaces (99-2); no anal-fin spines (100-0); three pelvic-fin elements (one spine and two reduced rays) (102-6); and

total number of vertebrae reduced to 20, which in our coding is

subsumed in the 26 or fewer category (93-0). See Tyler and

Sorbini (1996) for a smaller number of synapomorphies applicable to all fossil and extant tetraodontiforms and for those that

distinguish the most basal tetraodontiform clade of three Upper

Cretaceous families from all other, Eocene to present, families

of tetraodontiforms.

The most cogent analysis of zeiform characteristics previously available is that of Johnson and Patterson (1993), who

listed numerous characters as uniting all zeiforms. With most

of these we agree, but several others apply either at a different

level of universality or may form part of a larger transformation series, and several have convergently arisen in other

groups.

The synapomorphies listed by Johnson and Patterson (1993)

are as follows.

[I]. "Baudelot's ligament originating high on the exoccipital,

immediately under the vagus foramen." The condition of this

character (43) is unknown in half of the zeiform species studied

herein because of poor ligament visibility in some of the

cleared and stained material (or previous dissections in the vicinity of Baudelot's ligament having damaged it) and the disappearance of the ligament in most of the dry skeletal material. In

most of the zeiform taxa in which the determination can be

made, Baudelot's ligament originates on the first vertebra (43-

1), and our study indicates that it is this derived condition that

is ancestral for zeiforms (most outgroup taxa have the origin of

the ligament on the basioccipital, 43-0). Among zeiforms, only

Zeus and Zenopsis have a further derived condition of the ligament originating on the exoccipital (43-2). Thus, Baudelot's

ligament originating on the first vertebra is a synapomorphy for

zeiforms, but this character's support for the monophyly of zeiforms is somewhat questionable because of the large number of

taxa in which it presently is unknown.

We code Melamphaes 43-1 for this character, convergent

with the zeiform synapomorphy, because in both cases the origin of the ligament is on the first vertebra. The conditions

could be considered different, however, because in Melamphaes the origin is on the middle to lower part of the centrum

whereas in the zeiform ancestral condition it is higher and more

anterior, on an anterior process of the first vertebra just above

the posterior end of the basioccipital. In Zeus and Zenopsis the

origin has simply moved a little further forward and upward, to

a point on the exoccipital just above and in front of the anterior

process of the first vertebra.

[II]. "The dorsal fin endoskeleton has the distal portions of

the proximal-middle radials laterally expanded, and there are

no distal radials in the spinous portion except as tiny cartilaginous or partially ossified structures under the last few spines."

We find the degree of expansion of the distal portions of the

proximal-middle pterygiophore to be especially subjective and

difficult to define. The degree of distal expansion of the pterygiophores varies greatly in the outgroups, ranging from highly

expanded in Plectrypops and Sargocentron to not expanded in

Melamphaes. Within zeiforms, Cyttus and Parazen do not have

the distal end of the pterygiophores expanded, whereas oreosomatids have the distal end slightly expanded (the low, thin lateral flange along much of the length of the soft dorsal- and

anal-fin pterygiophores in Parazen forms a lateral prong distally, but we do not consider this comparable to a more general

expansion of the distal end of the pterygiophores). The remaining zeiforms do have distally expanded pterygiophores, even

though the degree of expansion varies. Because of the subjectivity in delineating character states, we did not include this

part of the character in our study.

The second part of this character, no distal radials in the

spinous portion except as tiny cartilaginous or partially ossified

structures under the last few spines (62-1), is found in all zeiforms, but it is also found in Parahollardia and all other tetraodontiforms. Johnson and Patterson (1993) did not include tetraodontiforms in their study, but this character would be a

synapomorphy for a putative clade of zeiforms and tetraodontiforms.

[III]. "The palatine has a specialized, mobile articulation

with the ectopterygoid, which is truncated dorsally, and the

metapterygoid is extremely reduced." We agree that the palatine articulation (18-1) is unique in zeiforms, and the reduced

metapterygoid (19-1) also is a synapomorphy of the group.

[IV]. "The anterior vertebral centra have an unusually flexible articulation, and ribs, when present, never occur anterior to

the fourth vertebra." The flexible articulation of the anterior

centra with the skull (41-1) has arisen convergently in Hoplostethus. It is true that ossified ribs (47) are nearly always absent from the first four vertebrae of zeiforms (the one exception

being Grammicolepis, where ribs begin on the third vertebra),

but our study indicates that the absence of all ossified ribs (47-3)

defines Cyttus, Cyttomimus, Macrurocyttus, and Parahollardia

and that the remaining zeiforms secondarily have ossified ribs

on at least some of the more posterior vertebrae; only members

of two of the most derived clades of zeiforms, the grammicole-
NUMBER 618 33

pidid Grammicolepis and the zeids Zeus + Zenopsis, have ossified ribs on most of the abdominal vertebrae, posterior to the

second and to the fourth vertebra, respectively.

[V]. "There is a distinctive configuration of the dorsal gill

arch elements, including upright columnar processes on PB2

and PB3, and absence of PB4 and UP4 (Rosen, 1973, figs. 101,

i(J2; 1984, figs. 22, 23)." We agree that most zeiforms have a

distinctive derived condition of upright columnar processes on

PB2 and PB3 (28, 29) and that all zeiforms have PB4 and UP4

absent (31-1). But there is much variation between zeiform

species in the length of the columnar processes on PB2 and

PB3, from relatively short or barely discernible and certainly

not prominent, as in Cyttus and a few other taxa (28-0, 29-0), to

especially long and prominent (28-2, 29-2). To distinguish

these different conditions, we use three codes for the length of

the processes: short or absent (state 0), moderately long (state

1), or long (state 2). Because the relatively short condition of

some taxa, like Cyttus and a few other zeiforms, is difficult to

distinguish from the short to essentially absent condition of the

processes in the outgroups (e.g., barely discernible or essentially absent in Capros compared with very short in Antigonia),

we code both the relatively short to absent conditions as 0.

Thus, our study does not recognize the columnar-process character as a synapomorphy of zeiforms, and only the moderately

long to long processes are phylogenetically informative for

some clades within zeiforms. A more detailed analysis than attempted here of the morphology of the upper pharyngeal bones,

including differences in how PB2 and PB3 are held together

and how both are held to the shaft-like suspensory PB1, will

surely yield a wealth of useful characters.

[VI]. "Below the frontals there is a continuous median cartilage extending between the ethmoid cartilage and pterosphenoids." We exclude this feature from our data base because we

could not determine it in many of the zeiform taxa (such as the

species represented in our materials only by dry skeletons or

poorly counter-stained specimens). We accept it, however, as a

highly probable zeiform synapomorphy because we find this

long cartilage well back into the orbit close to the pterosphenoids in Capromimus, Cyttomimus, Cyttus, Parazen, Xenolepidichthys, Zenion, Zenopsis, and Zeus, and it is absent in Antigonia, Capros, and Parahollardia.

[VII]. "In the caudal skeleton there is a full neural spine on

PU2, two epurals, and the parhypural is truncated proximally."

The presence of a full neural spine on PU2 (49-0) is not unique

to zeiforms in our data set. This condition occurs in Parahollardia and would be a synapomorphy for a putative clade of zeiforms and tetraodontiforms, as suggested by Rosen (1984). The

condition of two epurals in the caudal skeleton (51-1) does support a zeiform clade; however, many zeiforms have a further reduction to one epural (as does Parahollardia). The derived condition of the parhypural (52-1) supports a zeiform clade.

Thus, most of the many characters given by Johnson and

Patterson (1993) as uniting zeiforms do support the clade, some

consistently and some with homoplasy: palatine articulation;

reduced metapterygoid; two epurals or fewer; parhypural

slightly removed from and not embracing the centrum; probably some rendition of the PB2 and PB3 columnar process character when further studied, even though our coding may presently mask this possible synapomorphy; absence of PB4 and

UP4; flexible articulation of the anterior vertebrae; attachment

of Baudelot's ligament (although a different state is ancestral);

and median cartilage under the frontal in the orbit. Two others

support a putative clade of zeiforms and tetraodontiforms: no

spinous dorsal-fin distal radials and long NPU2.

In addition to these characters, the reduction in number of

supernumerary dorsal-fin spines from two to one has been suggested as a synapomorphy for zeiforms (Patterson, 1992; Tyler

and Sorbini, 1996; Tyler, 1998). Although there is a single supernumerary spine in all zeiforms (see Figures 6, 16, 24, 33,

39, 48, 57, 60, 70, 73, 74, 89, 90), compared with two in caproids and tetraodontiforms, this condition has arisen convergently among the outgroups studied herein in Plectrypops +

Sargocentron and in Melamphaes. Because this character is

highly variable among lower acanthomorphs, and this homoplasy is reflected among our outgroups, we did not include

it in our data set even though it clearly is a synapomorphy of

zeiforms.

Bannikov (1991), in disagreement with Rosen's (1984) contention that caproids were related to zeiforms and tetraodontiforms, argued for the close relationship of caproids with acanthuroids and the Eocene genus Acanthonemus (from Monte

Bolca, Italy) and the placement of caproids among Perciformes. The focus of Bannikov's noncladistic analysis was on

caproids and acanthuroids, and not on zeiforms, and we do not

further comment on it here because our own results concerning

caproid relationships are ambiguous.

Stiassny and Moore (1992) suggested characters that group

zeiforms (including caproids), scorpaeniforms, and perciforms

together in a group called "Higher Percomorpha." Their analysis was based on a single osteological system, albeit a highly

complex one, the anatomy of the pelvic girdle and its various

attachments, and it did not examine the majority of the taxa involved in the present study. Additionally, the results presented

were in the form of a majority-rule tree, an Adams consensus

tree (both of the preceding without topological constraints),

and selected trees. The only use of a strict consensus tree was

to summarize two (of the 154) most parsimonious trees, under

topological constraints to enforce monophyly of two taxa, one

of which was their zeiforms (i.e., zeiforms sensu stricto, as

used herein, plus caproids). Our strict consensus tree of all of

the 154 most parsimonious trees based on the data presented by

Stiassny and Moore (1992, table 1), and unconstrained topologically, resulted in a large polytomy with a single resolved

node. This node, with a branch support value of only one, supports a trichotomy of caproids (i.e., excluding zeiforms), scorpaeniforms, and perciforms.

Thus, although the finely detailed anatomical study of the

pelvis by Stiassny and Moore (1992) suggested characters sup-
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

porting a grouping of zeiforms and caproids with scorpaeniforms and perciforms, the analysis does not in fact support this

clade without the a priori constraints.

Two of the characters used by Stiassny and Moore (1992)

support a grouping of zeiforms and caproids: [VI] the presence

of an inter-pelvic ligament, which is also present in holocentrids, scorpaeniforms, and perciforms; and [VII] the absence of

an autogenous lateral pelvic radial in adults, which is shared

with at least atheriniforms, lampridids, scorpaeniforms, and

perciforms (but not holocentrids). A third character [VIII], the

shape of the pelvis in cross section, links caproids (i.e., excluding zeiforms), scorpaeniforms, and perciforms.

The present study, which mainly investigates relationships

within zeiforms, includes only the few features of the pelvic

girdle anatomy that differ between zeiform species. Few of

these features were used by Stiassny and Moore (1992), and we

omit several of the pelvic features used in their study of acanthomorph relationships. Because of the different taxa and anatomical features examined in our respective studies, and the

limited extent of the phylogenetic conclusions relative to zeiforms that can be drawn from Stiassny and Moore's (1992)

analysis, their conclusions were not useful in our attempts to

identify a closely related potential outgroup for zeiforms (as

used herein, excluding caproids).

Colgan et al. (2000) included two species (one represented

by three geographically different populations) of zeiforms

among a broad array of taxa in an analysis of basal acanthomorphs that used 12S rDNA and 16S rDNA and was based on

a minimum length, strict consensus tree of the combined data

set. Among the results were the Zeusfaber from Italy being the

sister group of the scombroid Rexea, and these two forming the

sister group of a monophyletic trichotomy consisting of Zeus

faber from Australia, Z.faber from Japan, and Cyttus austral is.

This strange aspect of that study is not further considered here.

Wiley et al. (2000) examined the relationships of lower acanthomorphs based on a maximum-parsimony, total-evidence

analysis of sections of the 12S mitochondrial ribosomal gene,

the 28S ribosomal nuclear gene, and 38 morphological transformation series. They found that Zeiformes formed the sister

group of Gadiformes (a relationship not previously proposed,

and one not tested in our study). This putative gadiform-zeiform relationship was recovered in only one of the three maximum parsimony trees generated for each data set individually

(Wiley et al., 2000, fig. 8). Unfortunately, the subsections of

this figure have been transposed, with the tree from the morphological data represented as fig. 8a, and not fig. 8c as the legend states (F. Santini, pers. comm., 15 Jan 2001). This has led

to minor discrepancies in the text. For example, they state (p.

344) that the morphological tree "did little better" than the

DNA data in playing a role in tree resolution, when in fact it resolves fewer nodes than either of the trees derived from the

DNA sets. It is clear to us, and to E.O. Wiley (pers. comm., 8

Feb 2001), that the possibility of gadiform-zeiform relationship

needs much further testing before giving it much credence.

Phylogenetic Summary

The monophyly of the zeiform fishes is very strongly supported in all four of the analyses, with branch support values of

10 for both of the unordered trees, with and without most meristic features (Figures 2, 4, respectively), and for the ordered

tree with meristics (Figure 1), and with a support value of nine

for the ordered tree without most meristics (Figure 3). The tree

statistics in all four of the analyses, however, indicate a moderate amount of homoplasy for the ordered trees (assuming the

trees do reflect the actual sequence of evolution of these taxa),

caused in part by our lack of accuracy in ordering the multistate

characters.

The four analyses of the data set produced trees that are different from one another, but many parts of the topology are

shared among all four trees. Three of the analyses (Figures

1-3) support Rosen's (1984) putative clade of caproids, zeiforms, and tetraodontiforms and also support a clade of zeiforms and tetraodontiforms; the fourth analysis (Figure 4),

which we contend is the best justified and most rational, leaves

these relationships unresolved.

In the unordered consensus tree of the reduced data set (Figure 4), the beryciforms (sensu Johnson and Patterson, 1993)

may be paraphyletic, with their relationships unresolved. The

only relationship resolved in the outgroups is the clade of

Plectrypops + Sargocentron.

Our hypothesis (Figure 4) of zeiform interrelationships neither supports nor rejects that of either Johnson and Patterson

(1993), in which zeiforms (excluding caproids) are the sister

group of beryciforms (excluding stephanoberyciforms) plus

percomorphs, or Rosen (1984), in which caproids are the sister

group to a tetraodontiform + zeiform clade. Three of the four

analyses (Figures 1-3), however, support Rosen's hypothesis,

even though the analysis that we propose as most rational does

not (Figure 4; the most conservative interpretation of the data,

with the fewest assumptions made).

Within zeiforms, the Cyttidae (Cyttoidei, genus Cyttus) is

the first group to split off the main branch, and it forms the sister group to the remaining zeiforms in three of the four analyses and has very strong support in all four analyses. The next

clade, which occurs in three of the four analyses, with weak to

good support, is the grouping of the remaining zeiforms (Zeioidei). Support for the Oreosomatidae varies from weak to very

strong among the four analyses. Within the Oreosomatidae,

Pseudocyttus is the sister group to the remaining genera

{Oreosoma + Neocyttus + Allocyttus) in all four of the analyses, but there is little agreement among the trees regarding the

relationships of the rest of this family. The Zeniontidae

(Zenion + {Capromimus + Cyttomimus)) has moderate to good

support in two of the four analyses. The Parazenidae (Parazen

+ (Cyttopsis + Stethopristes)) has weak to good support among

the four analyses. The Grammicolepididae (Macrurocyttus +

{Grammicolepis + Xenolepidichthys)) has moderate to strong

support among the four analyses. The Zeidae {Zeus + Zenopsis) has very strong support in all four of the analyses. In three
NUMBER 618 35

of the four analyses, Zeidae is the sister group to the Grammicolepidae.

In both of the two unordered analyses (Figures 2, 4), the zeiform families have the following phylogenetic sequence: Cyttidae, Oreosomatidae, Parazenidae, Zeniontidae, Grammicolepididae, and Zeidae. Familial relationships are mostly unresolved

in the ordered analysis of the reduced data set (Figure 3) and

are partially unresolved in that of the full data set (Figure 1),

with the latter having a different sequence of families than in

the two unordered analyses (Figures 2, 4).

SYSTEMATICS

Revised Familial Classification ofZeiformes

Of the five traditional zeiform families, only the Oreosomatidae is recognized herein as a monophyletic clade with the

usual grouping of genera: Allocyttus, Neocyttus, Oreosoma,

and Pseudocyttus. Two subfamilies are established for the

Oreosomatidae, with Pseudocyttus as the sister group of the

other three genera. The family-level concepts for the arrangement of all of the other genera must be moderately to radically

changed on the basis of the present study. The Grammicolepididae requires the least change: the inclusion for the first time of

Macrurocyttus along with Grammicolepis and Xenolepidichthys; two subfamilies are established, with Macrurocyttus as

the sister group of the other two genera. The formerly monotypic Zeniontidae becomes much expanded with the association for the first time of Zenion with Capromimus and Cyttomimus. The formerly monotypic Parazenidae becomes much

expanded with the association for the first time of Parazen with

Cyttopsis and Stethopristes; two subfamilies are established,

with Parazen as the sister group of the other two genera. The

Zeidae, formerly a large catch-all category, becomes restricted

to only Zeus and Zenopsis. A new monophyletic group, the

Cyttidae, is recognized for the three species of Cyttus, as the

sister group (Cyttoidei) of all other zeiforms (Zeioidei).

In the recognition of subfamilies, we take into account three

factors, listed in decreasing order of importance: the Bremer

support values for the monophyly of clades within a family,

based mostly on the analysis of the unordered reduced data set

(Figure 4) but with secondary consideration to the values in the

other three analyses (Figures 1-3); the degree of external and

internal morphological distinctiveness between these clades;

and the number of species within the clades. For example, with

a Bremer value of eight (very strong) in Figure 4 (and with values of 5, 8, and 12 in Figures 1-3) distinguishing the

Xenolepidichthys + Grammicolepis clade from Macrurocyttus

within the Grammicolepididae, and with substantial morphological differences between the two groups, there is good reason to recognize subfamilies therein, even though there are only

three species involved. Similarly, a Bremer value of three

(good) in Figure 4 (and values of 3, 5, and 8 in Figures 1-3) distinguishes the Cyttopsis + Stethopristes clade from Parazen

within the Parazenidae, and there are substantial morphological

differences between the two groups (supportive of subfamilial

recognition even though only three species are involved). More

subjective is the recognition of the Oreosoma+Neocyttus+Allocyttus clade as a subfamily distinct from Pseudocyttus within

the Oreosomatidae, because the Bremer value of this clade of

three genera is only two (moderate) in Figure 4 (and with values of 1, 2, and 4 in Figures 1-3). Nevertheless, we do so because there are four species within the Oreosoma + Neocyttus +

Allocyttus clade, and we presume that the several species of

Neocyttus not examined in this work will mostly fit the same

pattern as N. rhomboidalis. Thus, Oreosomatinae is one of the

most speciose subfamilies recognized herein, and we deem it

useful to distinguish this clade from the more morphologically

primitive Pseudocyttus. Within the Zeniontidae, we decline to

recognize the Capromimus + Cyttomimus clade as subfamilially

distinct from Zenion because the Bremer value of two (moderate) in Figure 4 does not receive additional support in the other

three analyses. The Bremer value for this clade is only one

(weak) in Figure 2 and two in Figures 1 and 3, and in the latter

two cases this clade is involved in multifamily polytomies.

Within the Zeidae, the Bremer values for the Zenopsis clade in

the four analyses range from moderate (2) to very strong (5),

but the species of this genus differ so little in overall morphology from those of Zeus that nothing useful would be gained in

recognizing these differences at other than the generic level.

The new information on the phylogenetic relationships of the

zeiform fishes presented here leads us to the following revised

classification of the group. Family-level names of various vintage are available for most of these categories, although usually

with a much different assemblage of genera than originally

used.

Order ZEIFORMES (as usually recognized, excluding caproids).—Dories

Suborder CYTTOIDEI (new)

Family CYTTIDAE Giinther, 1860 (as Cyttina for all zeiforms; newly restricted herein to Cyttus only).—Lookdown or bigeye dories

Genus Cyttus

Suborder ZEIOIDEI (new)

Family OREOSOMATIDAE Bleeker, 1859 (as Oreosomatoidei for Oreosoma only; used herein to include all of the

genera traditionally placed in that family).—Oreos or

warty dories

Subfamily PSEUDOCYTTINAE (new)

Genus Pseudocyttus

Subfamily OREOSOMATINAE Bleeker, 1859 (newly

recognized at the subfamilial level)

Genera Oreosoma, Neocyttus, and Allocyttus

Family PARAZENIDAE Greenwood et al., 1966 (originally

for Parazen only; used herein in a newly expanded

sense).—Slender dory and smooth dories

Subfamily PARAZENINAE Greenwood et al., 1966

(originally at the family level).—Slender dory

Genus Parazen
36

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Subfamily CYTTOPSINAE Greenwood et al., 1966

(originally Cyttopsidae, as a family-level synonym of Zeidae).—Smooth dories

Genera Cyttopsis and Stethopristes

Family ZENIONTIDAE Myers, 1960 (originally for Zenion

and its synonymous Cyttula; used herein in a newly expanded sense).—Armoreye or elongate dories (Zenion)

and rough-head or capro dories {Capromimus and Cyttomimus)

Genera Zenion, Capromimus, and Cyttomimus

Family GRAMMICOLEPIDIDAE Poey, 1873 (originally for

Grammicolepis only; used herein in a newly expanded

sense).—Tinselfishes or scaly dories, and dwarf dory

Subfamily MACRUROCYTTINAE Myers, 1960 (originally at the family level; used herein at the subfamilial level).—Dwarf dory

Genus Macrurocyttus

Subfamily GRAMMICOLEPIDINAE Poey, 1873 (newly

recognized at the subfamilial level).—Tinselfishes

or scaly dories

Genera Xenolepidichthys and Grammicolepis

Family ZEIDAE Latreille, 1825 (as the family Zeides, a

broad category for Zeus as well as Capros and several

other non-zeiform genera; used herein in a much restricted sense both from that and from present usage).—

Buckler dories, John dory, St. Peter fish

Genera Zeus and Zenopsis

Because the diagnostic characters that indicate the monophyly of the families and subfamilies of zeiforms are mostly internal osteological features, we include below characterizations

of each of these categories based mostly on external features

for use in systematic accounts, even though there is overlap in

many of these features between some of the families. We also

include a key to the families and subfamilies and a compilation

(Table 2) of meristic features for all of the taxa studied as an

aid to future studies on the systematics and phylogeny of zeiforms.

Characterizations of the

Families and Subfamilies ofZeiformes

CYTTIDAE (Figures 6-15).—Contained genus: Cyttus.

External: Dorsal-fin spines 8 or 9, rarely 10; dorsal-fin

rays 28-36; no locking mechanism between dorsal-fin spines.

Anal-fin spines 2, first spine sometimes immovable and fully

fused to distal end of pterygiophore, appearing as a broadbased, ventrally directed process from pterygiophore and tapering to a point distally; anal-fin rays 28-38; no locking mechanism between anal-fin spines. Pelvic fin 1,6; positioned under

pectoral-fin base. Pectoral-fin rays 11-18. Principal caudal-fin

rays 13; 2 or 3 procurrent rays above and below.

Scales on most of body small (l%-2% SL), rounded to

squarish, irregularly spinoid (C. australis, C. traversi) or relatively cycloid (C. novaezelandiae), sometimes deciduous (C

traversi); low sheath of scales along soft dorsal- and anal-fin

bases, none of these enlarged or much different in morphology

than surrounding scales. Scales along either side of ventral surface of isthmus slightly enlarged as scutes, strongly curved or

bent almost at 90° along middle of their width, forming a

somewhat flattened surface along much of distance between

ventral ends of cleithra to bases of pelvic-fin spines, many

scutes from right and left sides of isthmus meeting and slightly

overlapping along midline in a zipper-like alternation, with additional smaller spinoid scales often interspersed along area of

midline overlap; spiny processes at angle of curvature of right

and left zipper-like series of scutes sometimes (especially in C.

traversi) relatively regularly arranged one after the other to

form spiny, parallel lateral edges of the flat isthmus; in C. traversi the angular scutes right and left of bases of pelvic-fin

spines continue posteriorly, forming two scale rows separated

by a naked area enclosing anus, with the two rows converging

in front of anal-fin origin; small scute-like scales sometimes

(C traversi) present on dorsal profile from supraoccipital to

dorsal-fin origin. No buckler scales.

Mouth large, upper-jaw length (from midline front border of

premaxilla to ventral end of maxilla) somewhat greater than orbit diameter, and about equal to or slightly longer than snout

length. Ascending premaxillary process long, reaching behind

front border of orbit about one-half into orbit.

Body moderately deep to deep, about 1.2-2.0 times in SL.

No extended prejuvenile stage in early life history of taxa.

Internal: Total vertebrae 30-42 (usually 31 in C. australis,

32 in C. novaezelandiae, 41 in C. traversi); abdominal vertebrae 11; caudal peduncle vertebrae 4-7 (usually 4 or 5 in C.

australis and C. novaezelandiae, 6 or 7 in C. traversi). One or 2

vacant interneural spaces between space 6 and space 10; when

2 spaces vacant, vacancies separated by an occupied space (i.e.,

spaces are in 2 groups). Infraorbitals slender, 6-9 excluding

deep lachrymal. Epurals 2. Supraneural 1.

OREOSOMATIDAE (Figures 16-23).—Contained genera:

Pseudocyttus, Oreosoma, Neocyttus, Allocyttus.

External: Dorsal-fin spines 5-7, rarely 8; dorsal-fin rays

29-36; locking mechanism between first three dorsal-fin spines

in all genera except Pseudocyttus. Anal-fin spines 2 or 3, rarely

4; anal-fin rays 27-34; locking mechanism between first two

anal-fin spines. Pelvic fin usually 1,6 in all genera except 1,5 in

Pseudocyttus (rarely 1,7 in Oreosoma, sometimes 1,5 in Neocyttus); positioned under pectoral-fin base. Pectoral-fin rays

18-21, usually 19-21. Principal caudal-fin rays 13; 2 or 3

procurrent rays above and below.

Scales on most of body small (l%-2% SL), rounded to

squarish, usually spinoid, surface of many roughened with upright spinules (scales on sides of body in Oreosoma and

Pseudocyttus may be more cycloid and deciduous at some

stages (see Karrer, 1986), but all non-prejuvenile specimens

examined herein with many spinoid scales, especially toward

median-fin bases); low sheath of scales along soft dorsal- and

anal-fin bases, scales similar in morphology to and only
TABLE 2.—Some of the meristic characters of all of the zeiform species examined. Several of these species (denoted with "*") are not included in the analysis because they were examined only as radiographs

or limited dry skeletal material, and most of the characters used in the analyses are not available for them.

Most of the data are based on the materials examined for this work, but for some characters, especially

the number of dorsal- and anal-fin rays, we have incorporated data from the literature cited in the text. In

all cases, however, the most common condition(s) (underlined) is based on our materials. ("-" = not

applicable;"?" = unknown.)

Species

Cytlus no vaezelandiae

Cvttus auslralis

Cyttus traversi

Pseudocyttus maculatus

Oreosoma atlanticum

Neocyttus rhomboidalis

Allocyttus niger

Allocvttus folletti*

Parazen pacifwus

('ytlopsis roseus

Stelhopristes cos

Zcnion hololepis

Zenion kptolepis *

Capromimus abbreviates

Cyltomimus slclgis

Maerumcyttus acanthopodus

Xenolepidichthys dalgleishi

Grammkolepis brachiu.icuius

Zeus fahcr

Zeus capensis*

Zenapsis conchifer

Zenopsis nchulosus

Zcnopsis oblongus *

Dorsal-fin spines

8-9

8-9

9-10

5-6

6-8(7)

7

6 7

7-8

5-7(6=7)

8

7-8

6-7

6-7

6-7

7

7-8

5-7

5-6

6-7

9-10

9-10

8-10

8 9

8-9

Dorsal-fin rays

28-30

28-31

34-36

34-36

29-31

32-35

29 33

32-34

30-33

27-36

(30-32)

27-30

27-28

26-31

27-31

25-27

22-24

26-29

27-30

32-34

22-24

22-24

24-27

25-27

27-29

Anal-fin spines

2

1

2

2-

2-3

2-3

3-4

2 3

3

3

1

!

2

-2

1

2

2

2

2

0

2

2

4-5

4

3

3

3

Anal-fin rays

28-30

29-31

35-38

31-34

28-30

30-33

27 31

30-31

30-32

32-33

28-30

27-28

23-29

22-31

26-28

22-26

22-28

27-29

33-35

20-24

19-21

24-26

24-26

25-27

Pelvic-fin spines

0

0

0

1

I

1

1

1

1

1

1

1

0

0

0

Pelvic-fin rays

6

6

6

5

6-7

5-6

6

6

6

7

9

9

6

6

6

6

2-2

6

6-7

6-7

6

6

6

6

Pectoral-fin rays

11-12

11-12

16-18(17)

19-21(20)

19-21(20)

19-21(20)

18 20

19-20

19-21

15-16

13-15(14)

13-14

16-18

16-18

16-17

14-15

12-15(11)

14-15

14-15

12-15

(13-14)

13-14

12-13

11-12

11-13(12)

Caudal-fin

principal rays

13

13

13

13

13

13

13

13

13

13

13

13

13

13

13

13

12-16(13)

15

15

13

13

13

13

13

Caudal-finprocurrent rays

2

2

2-3

2-3

2

2-3

2

2-3

2

7-8

3-4

4

2-3

2-3

3-4

3

0

1

1

1

1

1

1

1

Total vertebrae

30-33(32)

30-32(31)

40-42(41)

42

38

38-41(40)

36 38(37)

40

40

34

32

32-33

27

27

32

28-30(29)

34-36

36-38(37)

43-46(44)

30-32(31)

30

34-36Q5J

34-36(31)

36

Abdominal

vertebrae

11

11

11

14-15

13-14

14-15

13 14

14

13

12

II

11

11

11

11

11

13

10

11

13-15(14)

14

14-15

14-Ji

14

Caudal pedunclevertebrae

4-5

4-5

6-7

6

7

6-7

5 7(6)

7

7

7

6

5-6

5

5

7

6-7

4

8

8-10

4-5

5

4-5

4

5

Total vacant

intemeural spaces

1

1

2

7

8

7-9(9)

6-9(7 8)

8

7

1

3^1

3

1-2

2

2

1

4-5

4

4

5

5

4-5

5

5

Position(s) of vacantintemeural spaces

6

6-8

7-8,9-10

2,4-9

2,4-9,8-11,12

2,4-11,11-12

2-12, extremely variable

2.4-9.10-11(2.4-9.11)

2,4-6,8,10-11

7

3,6-7,8-9

6-7.8-9.10(6-7.9)

5-6,7-8(6,8)

6-7,8(6,8)

5-6,8

7

3-5,6-9,9-10

2-6.4-7(3-5.7)

3-6.6-7.8(3-5,7)

4-5,6-8,8-10,11-12

(4.6-7.8-10.11)

4,6-7,8-10,11

4-5,6-8,9-10,11

(4-5,7-8,10)

4-5,7-8,10

3,5-6,8,10

Groups of vacant

intemeural spaces

-

-

1-2

2

3-4

2-3

2-4(2 3)

3

4

-

2-3

2-3

1-2

1-2

2

-

2-3

1-2

1-3(2)

3-4

3-4

2-1(3)

3

4

Infraorbitals

8-9

6-7

7

4

3

4

4

4

4

3

3

6-7

4

3

0

3-5(3^4)

5

5-8

•>

8-12

(9-10)

11-12

'7

Epurals

2

2

2

2

2

2

2

2

2

2

2

1

2

2

1

1

1

2

2

1

1

1

1

1

Supraneurals

1

1

1

1

1

1

1

1

1

0

1

1

1

1

1

1

1

1

1

0

0

0

0

0

'First anal-fin spine always fused to pterygiophore.

2

First anal-fin spine sometimes fused to pterygiophore.
38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

slightly if at all larger than surrounding scales. No specialized

bucklers or scutes.

Mouth large, upper-jaw length more or less equaling orbit diameter and much longer than snout length. Ascending premaxillary process moderate, reaching from front border of orbit to

behind front border and one-fifth into orbit.

Body moderately deep to deep, about 1.3-1.9 times in SL.

Extended pelagic prejuvenile stage to 100 mm or more SL,

with weak ossification, expanded abdominal region, much connective tissue in thick skin, and large conical protuberances

(hillocks) of transformed scales.

Internal: Total vertebrae 36-42; abdominal vertebrae

13-15; caudal peduncle vertebrae 6 or 7, rarely 5. Seven to 9,

rarely 6, vacant interneural spaces between spaces 2 and 12;

vacancies in 2-4 groups. Infraorbitals deep (with large pores,

bridges, or open lacunae between upper and lower edges), 3 or

4 excluding deep lachrymal. Epurals 2. Supraneural 1.

PSEUDOCYTTINAE.—Contained genus: Pseudocyttus. Pelvic fin 1,5; dorsal-fin spines usually 5, sometimes 6; first dorsal-fin spine long, about as long as or longer than second

spine; no locking mechanism between dorsal-fin spines;

lower border of dentary relatively smooth, without prominent serrations; no serrations along posterior border of supracleithrum; hypurals 1-4 fused and fused to centrum; total

vertebrae 42; opercles with delicate and relatively smooth

scales, without strong spiny ridges.

OREOSOMATINAE.—Contained genera: Oreosoma, Neocyttus, and Allocyttus. Pelvic fin usually 1,6 but sometimes

1,5 or 1,7 (see family account, above); dorsal-fin spines usually 6 or 7, sometimes 5 or 8; first dorsal-fin spine short,

much shorter than second spine; locking mechanism between

first three dorsal-fin spines; lower border of dentary with

multiple serrations behind symphysis; serrations along posterior border of supracleithrum; hypurals 1 and 2 fused and

fused to centrum, but hypurals 3 and 4, although fused, usually not fused to centrum; total vertebrae 36-41, usually

37-40; opercles armed with spiny scales along prominent radiating ridges.

PARAZENIDAE (Figures 24-38).—Contained genera: Parazen, Cyttopsis, Stethopristes.

External: Dorsal-fin spines 6-8; dorsal-fin rays 27-36

(usually 27-32); locking mechanism weakly developed between first two dorsal-fin spines in Parazen but absent in Cyttopsis and Stethopristes. One anal-fin spine, this sometimes

fused to pterygiophore in Cyttopsis; anal-fin rays 27-33 (first

anal-fin ray in Cyttopsis (second sequential element of anal fin)

loses its segmentation at about 100 mm SL and becomes more

ossified and spine-like in large specimens, and anal fin could be

considered to have two spines); no locking mechanism between

anal-fin spine and second element when latter becomes spinelike. Pelvic fin with no spine, rays 7 (Parazen) or 9 {Cyttopsis,

Stethopristes; Cyttopsis sometimes with 10 rays fide Heemstra,

1980); positioned either under or slightly in front of pectoralfin base (Cyttopsis, Stethopristes) or about midway between

anus and pectoral-fin base (Parazen). Pectoral-fin rays 13-16.

Principal caudal-fin rays 13; procurrent rays either 3 or 4 (Cyttopsis, Stethopristes) or 7 or 8 (Parazen) above and below.

Scales on most of body small (l%-2% SL), more or less

rounded, deciduous, either weakly spinoid (Parazen) or cycloid (Cyttopsis, Stethopristes); low sheath of scales along soft

dorsal- and anal-fin bases in Parazen, but scales absent along

these bases in Cyttopsis and Stethopristes (last two genera with

a firm, bony discontinuous ridge produced just below skin

along each side of soft dorsal- and anal-fin bases by strong lateral expansions at distal ends of pterygiophores, but this ridge

scaleless and not spiny). Large buckler scales (4%-7% SL)

present in Cyttopsis and Stethopristes (absent in Parazen)

along ventral midline of isthmus in front of bases of pelvic fins

and from pelvic fins to anus, many bucklers with a posteriorly

directed spiny process (sometimes also accessory smaller spiny

processes); bucklers continuous in a median keeled series in

Stethopristes but partially separated in Cyttopsis; about 7-12

bucklers in median series and sometimes fewer, less regularly

arranged additional smaller bucklers.

Mouth large, upper-jaw length slightly to much greater than

orbit diameter and snout length. Ascending premaxillary process moderate, reaching behind front border of orbit one-fifth

(Cyttopsis, Stethopristes) to one-third (Parazen) into orbit.

Body moderately deep to slender, about 1.3-2.5 (Cyttopsis,

Stethopristes) to 2.2-3.0 (Parazen) times in SL.

No extended prejuvenile stage in early life history of taxa.

Internal: Total vertebrae 32-34; abdominal vertebrae 11 or

12; caudal peduncle vertebrae 5-7 (usually 6 or 7). Vacant interneural spaces 1 (Parazen) or 3 or 4 (usually 3, Cyttopsis, Stethopristes), located between spaces 3 and 10 (usually 6 to 9);

when 3 or 4 spaces vacant, vacancies usually in 2 groups,

sometimes 3. Infraorbitals slender, 3 or 4 excluding deep lachrymal. Epurals 2 (Parazen, Cyttopsis) or 1 (Stethopristes). Supraneural 1 (Cyttopsis, Stethopristes) or 0 (Parazen).

PARAZENINAE.—Contained genus: Parazen. Dorsal-fin

spines 8; pelvic-fin rays 7; pelvic fin positioned about midway between pectoral-fin base and anus; pectoral-fin rays

usually 16; procurrent caudal-fin rays 7 or 8; scales spinoid;

no buckler scales; low sheath of scales along soft dorsal- and

anal-fin bases; total vertebrae 34; abdominal vertebrae 12;

caudal peduncle vertebrae 7; neural spines all oriented posterodorsally; no supraneural; 1 vacant interneural space; ventral

shaft of first dorsal-fin pterygiophore in middle of first interneural space, not contacting first vertebra; distal ends of dorsal- and anal-fin pterygiophores not much expanded (but

prongs present at upper end of thin lateral flanges along

pterygiophore); no lateral flange on lowermost pectoral-fin

radial; medial processes of basipterygia broadly overlapping

near pelvic-fin bases; 1 anal-fin pterygiophore anterior to first

haemal spine; 6 anal-fin pterygiophores anterior to third haemal spine; ossified epineurals on most abdominal vertebrae;

beryciform foramen represented by a groove along ceratohyal; hypurapophysis present. For additional differential con-
NUMBER 618 39

ditions between two subfamilies, see characters 15, 21, 29,

39, 53,57, and 71 in text, Appendix, Table 1, and Figures 4, 5.

CYTTOPSINAE.—Contained genera: Cyttopsis, Stethopristes. Dorsal-fin spines usually 6 (rarely 7) or 7 (rarely 8);

pelvic-fin rays 9; pelvic-fin positioned under or slightly in

front of pectoral-fin base; pectoral-fin rays usually 13 or 14

(rarely 15); procurrent caudal-fin rays 3 or 4; scales cycloid;

large buckler scales along ventral midline; no sheath of

scales along soft dorsal- and anal-fin bases; total vertebrae

usually 32; abdominal vertebrae 11; caudal peduncle vertebrae usually 6; some neural spines oriented anterodorsally or

at least vertically; 1 supraneural; usually 3 vacant interneural

spaces; ventral shaft of first dorsal-fin pterygiophore at front

of first interneural space, contacting neural arch and spine of

first vertebra; distal ends of dorsal- and anal-fin pterygiophores much expanded laterally; lateral flange on lowermost

pectoral-fin radial; no overlapping medial processes of basipterygia, halves of pelvis not in close contact along midline

near pelvic-fin bases; 2 anal-fin pterygiophores anterior to

first haemal spine; 4 anal-fin pterygiophores anterior to third

haemal spine; ossified epineurals absent, or present on a few

middle abdominal vertebrae; beryciform foramen present;

hypurapophysis absent. For additional differential conditions

between subfamilies, see character numbers given above under Parazeninae.

ZENIONTIDAE (Figures 39-59).—Contained genera: Zenion,

Capromimus, Cyttomimus.

External: Dorsal-fin spines 6-8; dorsal-fin rays 22-31;

locking mechanism between first three dorsal-fin spines {Zenion) or between second and third spines {Capromimus, Cyttomimus). Anal-fin spines 2 (second spine very short and not

externally visible in Zenion); anal-fin rays 22-31; locking

mechanism between anal-fin spines. Pelvic fin 1,6; positioned

under pectoral-fin base {Capromimus, Cyttomimus) or slightly

behind base {Zenion). Pectoral-fin rays 14-18 (usually 15 or

16). Principal caudal-fin rays 13; 2-4 procurrent rays above

and below.

Scales on most of body of moderate size (2%-4% SL),

rounded to squarish, spinoid; no sheath of scales along soft

dorsal- and anal-fin bases, but spiny processes present at angle

of scales bent around lateral expansions of distal ends of dorsal- and anal-fin pterygiophores, forming a spiny ridge along

each side of these fin bases. No specialized bucklers or scutes.

Mouth of moderate size to large, upper-jaw length greater

than snout length but either less than orbit diameter {Zenion) or

much greater than orbit diameter {Capromimus, Cyttomimus).

Ascending premaxillary process moderate, reaching behind

front border of orbit about one-fifth {Zenion) or about one-third

{Capromimus, Cyttomimus) into orbit.

Body moderately deep to relatively slender, about 2.0-3.0

times in SL.

No extended prejuvenile stage in early life history of taxa.

Internal: Total vertebrae 27-32; abdominal vertebrae 11;

caudal peduncle vertebrae 5-7. One or 2 vacant interneural

spaces between spaces 5 and 8; when 2 spaces vacant, vacancies usually separated by an occupied space. Infraorbitals deep

(with serrate vertical strengthening flanges, Zenion) or slender

{Capromimus, Cyttomimus), 3 or 4 {Capromimus, Cyttomimus)

or 6 or 7 {Zenion) excluding deep lachrymal. Epurals 1 {Capromimus, Cyttomimus) or 2 {Zenion). Supraneural 1.

GRAMMICOLEPIDIDAE (Figures 60-72).—Contained genera:

Macrurocyttus, Xenolepidichthys, Grammicolepis.

External: Dorsal-fin spines 5-7, usually 6 or 7; dorsal-fin

rays 26-34; locking mechanism between second and third dorsal-fin spines. Anal-fin spines 2 {Xenolepidichthys, Grammicolepis) or absent {Macrurocyttus); anal-fin rays 22-35; locking

mechanism between anal-fin spines only in Xenolepidichthys.

Pelvic fin either 1,6 or 1,7 (usually 1,6, Xenolepidichthys, Grammicolepis) or 1,2 or 1,3 (usually 1,3, Macrurocyttus); positioned

under or slightly in front of pectoral-fin base. Pectoral-fin rays

12-15, usually 13-15. Principal caudal-fin rays 15 {Xenolepidichthys, Grammicolepis) or 13 {Macrurocyttus, but highly variable, 12-16); procurrent rays above and below 1 {Xenolepidichthys, Grammicolepis) or 0 {Macrurocyttus).

Scales absent {Macrurocyttus, or highly deciduous and unknown) or narrow and greatly elongate vertically (to more than

25% SL) {Xenolepidichthys, Grammicolepis), with thin lateral

laminae curved posteriorly, relatively smooth edged {Xenolepidichthys) or with spinoid serrations {Grammicolepis); no

sheath of scales along soft dorsal- and anal-fin bases, but spiny

processes present {Xenolepidichthys, Grammicolepis) at angle

of scales bent around lateral expansions of distal ends of dorsal- and anal-fin pterygiophores, forming a spiny ridge along

each side of these fin bases. No specialized bucklers or scutes.

Mouth of moderate size, about equal to or slightly shorter

than orbit diameter and much longer than snout {Macrurocyttus), or small, about one-half or less of orbit diameter and much

shorter than snout {Xenolepidichthys, Grammicolepis). Ascending premaxillary process moderate, reaching from front

border of orbit to behind front border about one-fifth into orbit.

Body moderately deep to deep, about 1.1-2.6 times in SL.

No extended prejuvenile stage in early life history of taxa

(but Macrurocyttus bathypelagic and perhaps paedomorphic).

Internal: Total vertebrae 34-46; abdominal vertebrae

10-13; caudal peduncle vertebrae 4-10. Four or 5 vacant interneural spaces between spaces 2 and 10; vacancies in 1-3 (usually 2 or 3) groups. Infraorbitals absent {Macrurocyttus) or

slender, 3-5 excluding moderate lachrymal {Xenolepidichthys,

Grammicolepis). Epurals 1 or 2. Supraneural 1.

MACRUROCYTT1NAE.—Contained genus: Macrurocyttus.

Scales absent or highly deciduous and unknown; mouth

moderate in size, upper jaw much longer than snout, length

about equal to orbit diameter; principal caudal-fin rays usually 13; no procurrent caudal-fin rays; pelvic fin I, usually 3;

no pelvic-fin-ray serrations; no anal-fin spines; branchiostegal rays usually 2 + 4 = 6; postmaxillary process present

(short and broad); basisphenoid absent; total vertebrae usually 36; abdominal vertebrae 13; caudal peduncle vertebrae
40

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

4; all hypurals fused to centrum, hypural 5 not free; hypurapophysis absent; anterior end of parhypural simple, not

in contact with centrum; epural 1; symplectic without ventral

flange; medial processes of basipterygia broadly overlapping

near pelvic-fin bases; parietal absent; infraorbitals absent;

metapterygoid absent; ossified ribs absent; ossified epineurals absent; beryciform foramen represented by groove along

ceratohyal. For additional differential conditions between

two subfamilies, see characters 21, 37, 39, 58, 59, 69, and 99

in text, Appendix, Table 1, and Figures 4, 5.

GRAMMICOLEPIDINAE.—Contained genera: Xenolepidichthys, Grammicolepis. Scales greatly elongate vertically;

mouth small, upper-jaw length less than snout length and orbit diameter; principal caudal-fin rays 15; 1 procurrent caudal-fin ray above and below; pelvic fin I, usually 6; serrations on crests of several pelvic-fin rays; 2 anal-fin spines

(first spine greatly elongate in juveniles); branchiostegal rays

3+4 = 7; postmaxillary process absent; basisphenoid a short

shaft at front of roof of posterior myodome; total vertebrae

usually 37 or 44; abdominal vertebrae 10 or 11; caudal peduncle vertebrae 8-10; hypurals 1 through 4 fused to centrum but hypural 5 free; hypurapophysis present; anterior

end of parhypural with peg-and-socket-joint articulation with

centrum; epurals 2; symplectic with ventral flange; no overlapping medial processes of basipterygia, halves of pelvis

not in close contact along midline near pelvic-fin bases; parietal present; infraorbitals present; metapterygoid present; 1

to many ossified ribs present; many ossified epineurals

present; beryciform foramen present. For additional differential conditions between subfamilies, see character numbers

given above under Macrurocyttinae.

ZEIDAE (Figures 73-88).—Contained genera: Zeus, Zenopsis).

External: Dorsal-fin spines 8-10; dorsal-fin rays 22-29;

locking mechanism between first three dorsal-fin spines. Analfin spines usually 3 (Zenopsis) or 4 (Zeus), rarely 5 (Zeus);

anal-fin rays 19-27; locking mechanism between first two

anal-fin spines. Pelvic fin 1,6 or 1,7 (Zeus) or with no spine and

6 rays (Zenopsis, with first element in bilateral halves but unsegmented and thicker than other 5 rays); positioned slightly to

distinctly (especially in Zenopsis) in front of pectoral-fin base.

Pectoral-fin rays 11-15 (usually 12-14). Principal caudal-fin

rays 13; 1 procurrent ray above and below.

Scales on most of body small (l%-2% SL), cycloid, and embedded (Zeus) or absent entirely except for small lateral-line

scales (Zenopsis), but greatly enlarged buckler scales (up to

5%-l 1% SL, and many times larger than any normal scales on

body) present along bases of soft dorsal and anal fins, dorsal

series of bucklers extending anteriorly along much of base of

spinous dorsal fin in Zenopsis; these bucklers with a prominent, posteriorly curved spiny process, sometimes bifurcate

distally in some species (a smaller accessory spine present near

base of main spine in Zeus faber); bucklers along dorsal-fin

base about 4-14 and along anal-fin base about 4-11; other

bucklers present along ventral midline of isthmus in front of

bases of pelvic fins and from pelvic fins to anus, these series

usually a combination of single median bucklers and strongly

keeled (bent about 90°), paired bucklers from right and left

sides that slightly overlap along midline and often bear spiny

processes.

Mouth large, upper-jaw length much greater than orbit diameter and about equal to or slightly longer than snout length. Ascending premaxillary process moderate, reaching to about front

of orbit.

Body moderately deep to deep, about 1.3-2.2 times in SL.

No extended prejuvenile stage in early life history of taxa.

Internal: Total vertebrae 30-36; abdominal vertebrae

13-15 (usually 14 or 15); caudal peduncle vertebrae 4 or 5.

Four or 5 (usually 5) vacant intemeural spaces between spaces

3 and 12; vacancies in 2-4 (usually 3 or 4) groups. Infraorbitals

slender, 5-12 excluding moderate (Zeus) to slender (Zenopsis)

lachrymal. Epural 1. No supraneural.

Key to the Families and Subfamilies of Zeiformes

When two or more genera are included in a family or subfamily, keys are also provided

to the genera, with the exception of the three genera of Oreosomatinae. Although the

monophyly of the oreosomatins has substantial support, the phylogeny within the subfamily is only weakly supported, and the differences between Oreosoma and, especially, Neocyttus and Allocyttus are mostly either morphologically minor and internal or involve differences in scale patterns, allometrically variable proportions, and dorsal profile shapes

not thoroughly studied herein. The reader is referred to Heemstra (1980) and Karrer

(1986) for the best keys available to the genera of oreosomatids, and to Yearsley and Last

(1998) for the species of Neocyttus.

Large buckler scales with posteriorly directed, spiny (thorn-like) processes present

along bases of dorsal (4-14 scales) and anal (4-11 scales) fins; bucklers present or

absent along spinous dorsal-fin base; basal-plate length of larger bucklers ranging

from four times in orbit diameter to slightly longer than orbit diameter; bucklers

also along ventral midline from isthmus to anus ZEIDAE (B)

A.
NUMBER 618 41

BB.

Zeus faber

Large buckler scales along dorsal-fin base not extending forward beyond last dorsal-fin spine; anal-fin spines usually 4, rarely 5; anal-fin rays usually 19-24; dorsal-fin rays usually 22-24; first pelvic-fin element unbranched but stiff and

spine-like, followed by 6 or 7 branched rays (total pelvic-fin elements 7 or 8);

pectoral-fin rays usually 13 or 14, sometimes 12 or 15; scales on most of body

small, cycloid, embedded Zeus

Large buckler scales along dorsal-fin base extending forward under spinous dorsal

fin to about bases of third or fourth spines; anal-fin spines 3; anal-fin rays usually

24-27; dorsal-fin rays usually 24-29; first pelvic-fin element unbranched but

flexible and ray-like, followed by 5 branched rays (total pelvic-fin elements 6);

pectoral-fin rays usually 12, sometimes 11 or 13; scales absent on most of body,

except for those in lateral line Zenopsis

Zenopsis conchifer

ventral view of isthmus to anus

AA. No large buckler scales present along bases of dorsal and anal fins; bucklers present

or absent along ventral midline from isthmus to anus C

C. Scales along either side of ventral surface of isthmus slightly enlarged as scutes,

curved or bent almost 90° along middle of their width, forming a somewhat flattened surface along much of distance between ventral ends of cleithra to bases of

pelvic-fin spines, many scutes from right and left sides of isthmus meeting and

slightly overlapping along midline in a zipper-like alternation, with additional

smaller spinoid scales often interspersed along area of midline overlap; single

row of scutes on either side of a naked midline region between pelvic-fin bases

and anus or scutes absent; no buckler scales along ventral midline

CYTTIDAE, Cyttus

CC. No slightly enlarged scutes in a zipper-like arrangement along ventral surface of

isthmus (but large buckler scales present or absent along ventral midline from

isthmus to anus D

D. Pelvic fin with no spine, rays 7 or 9 PARAZENIDAE (E)
42

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Cyttus traversi above C. novaezelandiae

ventral view of isthmus to anus

Cyttus traversi

Dorsal-fin spines 8; pelvic-fin rays 7; pelvic-fin positioned about midway

between pectoral-fin base and anus; pectoral-fin rays usually 16, rarely

15; procurrent caudal-fin rays 7 or 8; scales spinoid; no large buckler

scales along ventral midline; low sheath of scales along soft dorsal- and

anal-fin bases PARAZENINAE, Parazen

Parazen pacificus

EE. Dorsal-fin spines usually 6 or 7, rarely 8; pelvic-fin rays 9; pelvic-fin positioned under or slightly in front of pectoral-fin base; pectoral-fin rays

usually 13 or 14, rarely 15; procurrent caudal-fin rays 3 or 4; scales cycloid; large buckler scales along ventral midline; no sheath of scales along

soft dorsal- and anal-fin bases CYTTOPSINAE (F)
NUMBER 618 43

Cyttopsis roseus

ventral view of isthmus to anus

DD.

Cyttopsis roseus

F. Dorsal-fin spines usually 7, sometimes 8; pectoral-fin rays usually 14,

sometimes 13 or 15; large buckler scales along ventral midline of

isthmus forming a partially separated spiny keel (a few bucklers not

overlapping, especially first, on isthmus, and second, between pelvic-fin bases); no honeycomb pattern on bones on top of head, opercles, and lachrymal-infraorbitals, only long ridges Cyttopsis

FF. Dorsal-fin spines usually 6, sometimes 7; pectoral-fin rays usually 13,

sometimes 14; large buckler scales along ventral midline of isthmus

forming a continuous, sharp-edged median spiny keel (nearly all

buckler scales strongly overlapping); honeycomb pattern on bones

on top of head visible externally Stethopristes

Pelvic fin with a spine, rays usually 6 or fewer (rarely 7) G

G Pectoral-fin rays 18-21, usually 19-21; low sheath of scales along soft

dorsal- and anal-fin bases, scales only slightly if at all larger than surrounding scales and of similar morphology; extended prejuvenile stage

to 100 mm or more SL, with weak ossification, expanded abdomen,

thick skin, and large conical protuberances of transformed scales

OREOSOMATIDAE (H)

H. Pelvic fin 1,5; dorsal-fin spines usually 5, sometimes 6; first dorsal-fin

spine long, about as long as or longer than second spine; no locking

mechanism between dorsal-fin spines

PSEUDOCYTTirsAE, Pseudocyttus
44

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

HH. Pelvic fin usually 1,6 but sometimes 1,5 or 1,7; dorsal-fin spines usually

6 or 7, sometimes 5 or 8; first dorsal-fin spine short, much shorter

than second spine; locking mechanism between first three dorsal-fin

spines OREOSOMATINAE, Oreosoma, Neocyttus, AUocyttus

Neocyttus rhomboidalis

GG Pectoral-fin rays 12-18, usually 13-16; no sheath of scales along soft

dorsal- and anal-fin bases, but, except one species in which scales are

absent, spiny processes present at angle of scales bent around lateral expansions of distal ends of dorsal- and anal-fin pterygiophores, these processes forming spiny ridge along each side of fin bases; no extended

prejuvenile stage I

I. Scales on most of body of moderate size (2%-4% SL), rounded to

squarish, spinoid; procurrent caudal-fin rays 2-4

ZENIONTIDAE (J)

J. Dorsal-fin spines usually 6, sometimes 7; locking mechanism between first, second, and third dorsal-fin spines; second anal-fin

spine below surface of skin and not visible externally; pelvic fin

positioned slightly behind pectoral-fin base; upper-jaw length less

than orbit diameter; infraorbital bones very deep, covering most

of cheek below orbit, with serrate vertical strengthening flanges;

no anteromedial processes on medial (lower) surfaces of pelvicfin rays Zenion

Zenion hololepis

JJ. Dorsal-fin spines usually 7 or 8; locking mechanism between only

second and third dorsal-fin spines; second anal-fin spine short,

about equal to length of first spine or slightly less than one-half its

length, but visible externally; pelvic fin positioned under pectoralfin base; upper-jaw length greater than orbit diameter; infraorbital

bones slender and tubular, not covering most of cheek below or-
NUMBER 618 45

bit; anteromedial processes present as broad flanges along basal

regions of pelvic-fin rays K

K. Dorsal-fin spines usually 7; dorsal-fin rays usually 25-27; analfin rays 26-28; pectoral-fin rays usually 16, sometimes 17; no

prominent posterior flange at upper end of postcleithrum; no

ectopterygoid teeth; serrations present on pelvic-fin-ray

flanges Capromimus

KK. Dorsal-fin spines usually 8, sometimes 7; dorsal-fin rays usually

22-24; anal-fin rays 22-26; pectoral-fin rays usually 15, sometimes 14; prominent posterior flange at upper end of postcleithrum, flared posterolaterally just behind posterior border of

opercular flap; ectopterygoid teeth present; no serrations on

pelvic-fin-ray flanges Cyttomimus

Cyttomimus stelgis

Scales narrow and greatly elongate vertically (to more than 25% SL)

on most of body or absent; procurrent caudal-fin rays 1 or 0

GRAMMICOLEPIDIDAE (L)

L. Scales absent (or highly deciduous and unknown); mouth moderate

in size, upper jaw much longer than snout and length about equal

to orbit diameter; principal caudal-fin rays usually 13; procurrent

caudal-fin rays 0; pelvic fin with spine and usually 3 rays; anal-fin

spines 0; branchiostegal rays usually 2 + 4 = 6

MACRUROCYTTINAE, Macrurocyttus

Macrurocyttus acanthopodus

LL. Scales greatly elongate vertically; mouth small, upper-jaw length

less than snout length and orbit diameter; principal caudal-fin rays

15; 1 procurrent caudal-fin ray above and below; pelvic fin with

spine and usually 6 rays; anal-fin spines 2; branchiostegal rays 3 +

4 = 7 GRAMMICOLEPIDINAE (M)
46

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

M. Dorsal-fin spines usually 6, sometimes 5; dorsal-fin rays usually

27-30; anal-fin rays usually 27-29; pectoral-fin rays usually

14, sometimes 15; locking mechanism between anal-fin spines;

second anal-fin spine short, less than one-half length of first

spine in adults (first spine greatly elongate in juveniles); elongate vertical scales with relatively smooth edges; no prominent

horizontal crests on elongate scales and protruding laterally

from body at any size Xenolepidichthys

Xenolepidichthys dalgleishi

MM. Dorsal-fin spines usually 7, sometimes 6; dorsal-fin rays 32-34;

anal-fin rays usually 33-35; pectoral-fin rays usually 15, sometimes 14; no locking mechanism between anal-fin spines; second anal-fin spine long, more than one-half length of first spine

in adults (first spine greatly elongate in juveniles); elongate vertical scales with serrations; in juveniles and young adults up to

about 10 horizontal scale crests protrude prominently from lateral surface of body, crests formed from lateral projections

along middle regions of 2 to 5 adjacent scales, diminishing in

size in large adults (faint remnants of these crests still present as

thickened, low emarginations in a 285 mm SL specimen)

Grammicolepis

Conclusion

The present study elucidates the phylogeny of the zeiform

fishes and leads to a much revised systematic arrangement of

the genera into newly recognized monophyletic families and

subfamilies. However, this is but a first effort at understanding

zeiform phylogeny, and refinements can be expected when the

several species not included herein are taken into consideration

and, especially, when data sets based on molecular genetics,

soft anatomy (e.g., muscles and nerves), and other non-osteological features are used. We hope that the present study, based

mostly on bony structures, will be an encouragement to such

further analyses.
Appendix: Character-State Transformations for the

Unordered Reduced Data Set Consensus Tree

(Letters correspond to branches in Figure 4; numbers correspond to the characters and

their states used herein. The meristic characters not included in the analysis (93-103)

resulting in Figure 4 were optimized onto the tree and are shown in Figure 5.)

A: 2-2,6-1,13-1,16-1,20-1,27-1,43-1,53-4,54-1,59-0,63- Y:

0, 65-0,70-1,71 -0, 72-1,75-0,87-1 Z:

B: 5-1, 10-2, 13-1, 21-0, 22-2, 25-2, 46-0, 48-3, 54-1, 59-0, A1:

64-1, 65-0, 69-0, 70-3, 71-0, 76-0, 80-4, 82-1, 84-1, 85-2,

90-1 B1:

C: 10-2,25-1,31-1,41-1,43-3,46-0,48-1,63-0,64-2,70-3, C:

72-6, 83-2, 84-2, 88-4

D: 3-0, 13-1, 25-1, 35-0, 38-1, 59-1, 63-0, 72-3, 75-0, 77-1

E: 3-0, 4-1, 6-1, 10-0, 27-1, 33-3, 36-2, 38-1, 59-1, 63-0, D1:

69-2, 71-2, 72-2, 75-0, 80-1, 81-1

F: 3-0, 8-1, 25-2, 33-1, 35-0, 63-0, 64-2, 70-0, 71-0, 72-0, E':

75-0,81-1

G: 22-2,77-1 P:

H: 6-1, 10-0, 17-0, 20-1, 38-1, 60-1, 61-0 G':

I: 3-0, 4-1, 11-1, 15-3, 21-3, 22-2, 33-3, 34-1, 37-1, 38-1,40-

1,47-4, 60-1, 64-0, 69-2, 70-4, 71-2, 72-5, 77-1, 80-2, 84-1 FT:

J: 4-1, 9-2, 10-0, 12-1, 16-1, 27-1, 36-2, 37-1, 38-1, 40-1,

44-1, 60-1, 64-2, 69-2, 70-5, 71-3, 72-7, 77-1, 80-1, 81-1, F:

84-1

K: 1-1, 2-2, 4-1, 6-1, 9-0, 11-2, 12-1, 13-1, 16-1, 25-1, 27-1, J1:

33-2, 34-1, 37-1, 39-0, 40-1, 47-3, 49-0, 50-1, 51-2, 55-1,

59-1, 60-1, 62-1, 64-0, 69-3, 70-1, 72-0, 75-0, 80-3, 85-3, K':

86-2 L1:

L: 3-0, 10-0, 16-1, 18-1, 19-1, 21-1, 23-1, 24-1, 25-2, 26-0,

31-1, 35-0, 37-1, 40-1, 41-1, 43-1, 44-1, 49-0, 50-1, 51-1, M':

52-1, 53-2, 54-1, 55-1, 60-1, 62-1, 64-1, 73-1 (monophyly

ofZeiformes) N':

M: 9-3, 15-2, 22-2, 30-1, 45-1, 46-0, 47-3, 48-1, 74-1, 83-1,

89-1 (monophyly of Cyttus, Cyttidae, Cyttoidei) O':

N: 22-1, 67-1, 87-1 (autapomorphies of Cyttus novaezelandiae) P':

O: 77-1 (monophyly of Cyttus australis + C. traversi)

P: 36-1,91-1 (autapomorphies of Cyttus australis) Q':

Q: 50-0, 67-2, 72-2 (autapomorphies of Cyttus traversi)

R: 28-2, 29-1,47-1, 66-1 (monophyly of Zeioidei) R1:

S: 10-1, 28-1, 59-1, 92-1 (monophyly of Oreosomatidae)

T: 3-1 (autapomorphy ofPseudocyttus maculatus) S':

U: 22-2, 36-1, 53-1, 57-3, 77-1 (monophyly of Oreosomatinae)

V: 3-1, 28-0, 29-0, 36-0, 48-1,65-0,71 -0 (autapomorphies of T:

Oreosoma atlanticum) U':

W: (no autapomorphies for Neocyttus rhomboidalis)

X: 33-1,58-2, 72-2 (monophyly of Allocyttus verrucosus + A. V:

niger) W1:

(no autapomorphies for Allocyttus verrucosus)

28-2, 29-2, 53-2 (autapomorphies of Allocyttus niger)

46-0, 72-3, 85-2 (monophyly of unnamed clade encompassing taxa Parazen through Zenopsis in Figure 4)

45-2, 75-0, 82-1, 83-2 (monophyly of Parazenidae)

15-1, 21-2, 29-2, 33-1, 39-0, 44-0, 50-0, 53-1, 57-1, 61-0,

64-0, 69-2, 71-2, 81-0, 85-0 (autapomorphies of Parazen

pacificus)

48-3,72-1, 74-1, 87-1, 88-1, 90-1 (monophyly of Cyttopsinae)

22-1, 28-1, 48-2, 67-1 (autapomorphies of Cyttopsis roseus)

5-1, 37-0, 51-2, 70-1 (autapomorphies of Stethopristes eos)

51-2, 57-3, 76-0, 90-2 (monophyly of unnamed clade encompassing taxa Zenion through Zenopsis in Figure 4)

14-1,22-2,33-2, 37-0, 68-1, 77-1, 85-0 (monophyly of Zeniontidae)

9-2, 10-2, 20-1, 21-2, 38-1, 42-1,44-0, 51-1, 81-1 (autapomorphies of Zenion hololepis)

15-1, 47-3, 57-2, 71-2, 72-4, 83-2 (monophyly of

Capromimus + Cyttomimus)

47-2, 84-2 (autapomorphies of Capromimus abbreviatus)

14-0, 17-0, 50-0, 68-0, 76-1 (autapomorphies of Cyttomimus stelgis)

2-2, 8-1, 58-2, 87-3 (monophyly of unnamed clade encompassing taxa Macrurocyttus through Zenopsis in Figure 4)

3-1, 28-0, 29-0, 32-1, 35-1, 42-1, 46-1, 57-2 (monophyly

of Grammicolepididae)

1-1, 9-0, 19-2, 33-1, 37-0, 39-0,47-3,48-3, 53-5, 69-2, 72-

2, 85-0 (autapomorphies of Macrurocyttus acanthopodus)

2-1, 12-1, 13-1, 20-1, 21-0, 30-1, 50-0, 51-1, 52-2, 58-1,

59-1, 84-1, 87-2 (monophyly of Grammicolepidinae)

46-0, 65-0, 68-1 (autapomorphies of Xenolepidichthys dalgleishi)

9-2, 22-2, 47-5, 66-0 (autapomorphies of Grammicolepis

brachiusculus)

6-1,7-1, 9-5, 11-1, 14-2, 36-2, 43-2, 47-0, 48-1, 56-1, 59-

3, 64-0, 78-1, 79-1, 80-1, 83-1, 86-1, 88-3, 90-1 (monophyly of Zeidae)

2-1, 36-1, 58-3, 87-1, 88-2 (autapomorphies of Zeus faber)

8-2, 39-0, 82-1 (monophyly of Zenopsis conchifer + Z.

nebulosus)

9-4, 48-0, 91-2 (autapomorphies of Zenopsis conchifer)

72-4 (autapomorphy of Zenopsis nebulosus)

41
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Figures
FIGURE 6.—Cyttidae. Skeleton of

Cytlus traversi, AMNH 92052,

215 mm SL, lateral view. Character-state (for selected unlabeled

features not indicated in detailed

osteologieal views for species of

the family): 8-0 deep lachrymal;

10-0 slender, tubular infraorbitals;

44-1 some neural spines oriented

anterodorsally; 45-1 vacuities of

moderate size in abdominal haemal arches; 46-0 abdominal haemal spines with prominent ventral

processes; 47-3 no ossified ribs;

48-1 ossified epineurals on only a

few anterior abdominal vertebrae;

57-0 no dorsal-fin spine locking

mechanism; 58-1 two groups of

vacant interneural spaces; 59-2

three dorsal-fin pterygiophores

anterior to fourth neural spine; 61-

1 ventral end of shaft of first dorsal-fin pterygiophore at front of

first interneual/prcneural

space, contacting skull and first

vertebra between the two sides of

first neural arch and spine; 63-1

symmetrical pterygiophores; 64-1

one supraneural; 67-2 fusion of

first anal-fin spine to pterygiophore in all specimens; 73-1 dorsal-, anal-, and pectoral-fin rays

unbranched; 75-1 single postcleithrum; 78-0 ventral end of suprucleithrum simple; 81-2 pelvic fin

under pectoral fin; 99-2 two vacant

interneural spaces.
NUMBER6I8 53

19-1

FIGURE 7.—Cyttidae. Skull and first vertebra of Cyttus traversi, AMNH 92029,

316 mm SL, lateral view. The lachrymal, infraorbitals, and nasal are removed,

and the articulations between some of the otic-occipital bones are not shown

because they are unclear in this dry skeleton. On the dentary, parallel lines show

the more anterior and shorter of the two dentary cartilages, overlapping the

wavy lines of the more posterior cartilage. Character-state: 2-0 basisphenoid a

long shaft; 6-0 supraocular serrations on frontal; 15-2 ascending premaxillary

process reaching back about one-half into orbit; 18-1 palatine articulation pivotal; 19-1 reduced metapterygoid; 20-0 no symplectic ventral flange; 21-1 two

dental cartilages of moderate length, the first shorter; 22-2 multiple dentary serrations; 40-1 first neural arch and spine closely applied to skull.

20-0

22-2

FIGURE 8.—Cyttidae. Suspensorium of Cyttus austratis, AMS 1.32386001, 60

mm SL, lateral view. Wavy lines between the base of the lateral ethmoid and

the base of the palatine and the anterodorsal region of the mesopterygoid show

ligaments; small circles in mechanical stippling show the cartilages at the

bases of the palatine and lateral ethmoid. The dorsal end of the ectopterygoid

and the anterodorsal region of the mesopterygoid are especially sturdy for support of the joint with the ventral end of the palatine. Character-state: 18-1

palatine articulation pivotal; 19-1 reduced metapterygoid; 20-0 no symplectic

ventral flange.
54

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 9.—Cyttidae and other families. Basihyal and the three basibranchials (anterior ends downward), dorsal

view: A, Zeusfaber (Zeidae), USNM 320014, 89 mm SL; B, Cyttopsis roseus (Parazenidae), AMNH 29460, 75

mm SL; C, Stethopristes eos (Parazenidae), USNM 226570, 91 mm SL; D, Cyttus australis (Cyttidae), AMS

1.32386001, 60 mm SL. From bottom to top, the four elements in the series are the basihyal and the first, second,

and third basibranchials. Character-state: 30-0 first basibranchial not depressed below surface of basihyal (A-C);

30-1 first basibranchial depressed below surface of basihyal. (All scale bars=2 mm.)

FIGURE 10.—Cyttidae. Pelvis, dorsal view (anterior to left): A, Cyttus traversi, AMNH 99777, 455 mm SL; B,

Cyttus australis, AMNH 91908, 302 mm SL. Character-state: 85-1 basipterygia in contact in midline but not

overlapped; 86-0 short posterior process of pelvis. (Scale bars=5 mm.)
NUMBER 618

55

36-0

37-1

33-0 36-1

38-0

FIGURE 11.—Cyttidae and Zeniontidae. Hyoid arch, lateral (A, B) and medial (C) views: A, Cyttus traversi (Cyttidae), AMNH 92029, 316 mm SL; B, Cyttus australis, AMS 1.32386001, 60 mm SL; C, Capromimus abbreviates

(Zeniontidae), AMS 1.23650001, 61 mm SL. Character-state: 33-0 beryciform foramen present; 33-2 beryciform

foramen represented by deep concavity on dorsal surface of ceratohyal; 34-0 branchiostegals spread over/along

both epihyal and ceratohyal (B, C; would apply to A if branchiostegals were shown); 35-0 ceratohyal notches

present (A-C); 36-0 ceratohyal-epihyal articulation through cartilage only; 36-1 ceratohyal-epihyal articulation

either through cartilage (this specimen) or becoming interdigitated in larger specimens; 37-0 epihyal about same

depth as rear of ceratohyal; 37-1 epihyal less deep than rear of ceratohyal (A, B); 38-0 urohyal no longer than ceratohyal (B, C; would apply to A if urohyal were shown).
56 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

102-0

FIGURE 12.—Cyttidae and other families.

Basal region of the right pelvic fin, ventral

view: A, Stethopristes eos (Parazenidae),

USNM 226570, 91 mm SL; B, Cyttus australis (Cyttidae), AMS I.32386OO1, 60 mm SL;

C, Capromimus abbreviatus (Zeniontidae),

AMS 1.23650001, 61 mm SL. Characterstate: 82-0 pelvic-fin spine present (B, c); 82-

1 no pelvic-fin spine; 83-1 anterolateral processes as prongs from bases of rays; 83-2

anterolateral processes as broad flanges along

basal region of rays (A, C); 84-2 serrations on

broad flanges (C); 102-0 total of nine pelvicfin elements; 102-2 total of seven pelvic-fin

elements.

102-2

84-2
NUMBER (>18 57

FIGURE 13.—Cyttidae and other families. Right postcleithrum and adjacent region of the cleithrum, medial

view: A, Cyttopsis roseus (Parazenidae), USNM 187873, 72 mm SL; B, Stethopristes eos (Parazenidae), USNM

226570, 91 mm SL; C, Zenopsis conchifer (Zeidae), FMNH 67090, 58 mm SL; D, Cyttus traversi (Cyttidae),

AMNH 92029, 316 mm SL; E, Cyttus australis. AMNH 91932, 250 mm SL; F, Cyttomimus stelgis (Zeniontidae), USNM 306156, 76 mm SL. Character-state: 75-0 postcleithrum in two pieces; 75-1 single postcleithrum

(C-F); 76-0 no flange on single postcleithrum (C); 76-1 flange on single postcleithrum (D-F); 79-1 cleithral process present (C).
58

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

74-1

FIGURE 14.—Cyttidae and other families. Lowermost pectoral-fin radial, lateral view: A, Stethophstes eos (Parazenidae), USNM 226570, 91 mm SL; B, Cyttus australis (Cyttidae), AMS 1.32386001, 60 mm SL; C, Cyttopsis

roseus (Parazenidae), AMNH 29460, 74 mm SL; D, Zenopsis conchifer (Zeidae), CAS 47401, 54 mm SL; E,

Zeusfaber (Zeidae), USNM 320014, 89 mm SL; F, Capromimus abbreviatus (Zeniontidae), AMS 1.23650001,

61 mm SL. Character-state: 74-0 no lateral flange; 74-1 lateral flange present. (All scale bars=l mm.)
NUMBER 618

59

49-0

50-0

FIGURE 15.—Cyttidae. Caudal skeleton, lateral view: A, Cyttus australis, AMS

1.32386001, 60 mm SL; B, Cyttus traversi, AMNH 99778, 445 mm SL. Character-state: 39-1 PU3 first modified vertebra in caudal peduncle (A, B); 49-0

NPU2 long (A, B); 50-0 hypurapophysis present; 50-1 no hypurapophysis; 51-1

two epurals (A, B); 52-1 anterior end of parhypural slightly removed from and

not embracing centrum (A, B); 53-2 hypurals 1 + 2 + 3 + 4 fused and fused to

centrum (A, B); 54-1 no uroneural (A, B); 55-1 no stegural (A, B).
FIGURE 16.—Oreosomatidae.

Skeleton of Neocyttus rhomhoidalis, AMNH 91746, 232 mm SL,

lateral view. Character-state (for

selected features not indicated in

detailed osteological views for

species of the family): 8-0 deep

lachrymal; 10-1 deep infraorbitals

with large pores and bridges; 15-0

ascending premaxillary process

reaching no further than front of

lateral ethmoid and orbit; 44-1

some neural spines oriented

anterodorsally; 45-0 no vacuities

in abdominal haemal spines; 46-1

abdominal haemal spines without

prominent ventral processes; 47-1

ossified ribs only on last few

abdominal vertebrae; 48-0 ossified

epineurals on most abdominal vertebrae; 57-3 locking mechanism

between first, second, and third

dorsal-fin spines; 58-1 two groups

of vacant interneural spaces; 59-1

two dorsal-fin pterygiophores

anterior to fourth neural spine; 63-

1 symmetrical pterygiophores; 66-

1 locking mechanism between first

and second anal-fin spines; 73-1

dorsal-, anal-, and pectoral-fin rays

unbranched; 75-1 single postcleithrum; 78-0 ventral end of supracleithrum simple; 81-2 pelvic fin

under pectoral fin; 99-8 eight

vacant interneural spaces (minority condition; usually nine spaces

vacant, 99-9). Most specimens

have three anal-fin spines (100-3);

four in this specimen (100-4).
NUMBER 618 61

1-0

60-1

FIGURE 17.—Oreosomatidae. Skull and first vertebra ofNeocyttus rhomboidalis, AMNH 91743, 295 mm SL, lateral view (some details, especially of the first vertebra, from AMNH 93612,243 mm SL). Two obliquely oriented

shafts have only their basal regions indicated: the anterior shaft is the base of the supraneural and the posterior

shaft is the base of the first dorsal-fin pterygiophore. Character-state: 1-0 parietal present; 2-0 basisphenoid a long

shaft; 3-0 vomerine teeth present; 11-0 dermosphenotic a distinctly separate ossification; 40-1 first neural arch

and spine closely applied to skull; 42-0 no long dorsal extension of first neural spine above attachment to skull;

48-0 ossified epineurals on most abdominal vertebrae (only the first of the long series, on the first vertebra,

shown); 60-1 ventral end of shaft of first dorsal-fin pterygiophore in first intemeural/preneural space; 61-1 ventral

end of shaft of first dorsal-fin pterygiophore at front of first intemeural/preneural space, contacting skull and first

vertebra between the two sides of first neural arch and spine; 64-1 one supraneural; 80-0 one long extrascapular.
62

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

3-0

FIGURE 18.—Oreosomatidae. Skull and first vertebra of Neocyttus rhomboidalis, AMNH 91743, 295 mm SL,

dorsal (left) and ventral views (some details, especially of the first vertebra, from AMNH 93612, 243 mm SL).

Character-state: 1-0 parietal present; 3-0 vomerine teeth present; 4-0 no parasphenoid opening into posterior

myodome; 6-0 supraocular serrations on frontal; 11-0 dermosphenotic a distinctly separate ossification; 40-1 first

neural arch and spine closely applied to skull; 80-0 one long extrascapular.
NUMBER 618

63

20-0

FIGURE 19.—Oreosomatidae. Suspensorium and opercular bones of Neocyttus rhomboidalis, AMNH 93612, 243

mm SL, lateral view. The adherent spiny scales on the subopercle and the lower region of the opercle are not

shown. Character-state: 18-1 palatine articulation pivotal; 19-1 reduced metapterygoid; 20-0 no symplectic ventral flange.

38-0

35-0

FIGURE 20.—Oreosomatidae. Hyoid arch and branchiostegal rays of Neocyttus rhomboidalis, AMNH 93613.

255 mm SL, lateral view. Character-state: 33-0 beryciform foramen present; 34-0 branchiostegals spread

over/along both epihyal and ceratohyal; 35-0 ceratohyal notches present; 36-1 ceratohyal-epihyal articulation

either through cartilage (this specimen) or becoming interdigitated in larger specimens; 37-1 epihyal less deep

than rear of ceratohyal; 38-0 urohyal no longer than ceratohyal.
64 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 21.—Oreosomatidae. Pelvis ofNeocyttus rhomboidalis, AMNH 93612, 243 mm SL, dorsal view. Character-state: 85-1 basipterygia in contact in midline but not overlapped; 86-0 short posterior process of pelvis.

FIGURE 22.—Oreosomatidae. Right cleithrum

and adjacent region of the single postcleithrum of Neocyttus rhomboidalis, AMNH

91743, 295 mm SL, medial view. Characterstate: 75-1 single postcleithrum; 76-1 flange

on single postcleithrum.

76-1
NUMBER 618 65

51-1

54-1

55-1

49-0 51-2

97-2

FIGURE 23.—Oreosomatidae. Caudal skeleton of Neocyttus rhomboidalis, lateral view: A, AMNH 93613, 255

mm SL; B, AMNH 91746, 232 mm SL. The lengths of HPU3 are different in the two specimens; A has the more

normal configuration; in B, PU2 may be a compound vertebra, as indicated by deep grooves along the lengths of

the neural and haemal spines. Character-state: 39-1 PU3 first modified vertebra in caudal peduncle; 49-0 NPU2

long; 50-1 no hypurapophysis; 51-1 two epurals; 51-2 one epural (minority condition); 52-1 anterior end of parhypural slightly removed from and not embracing centrum; 53-1 hypurals 1 + 2 fused and fused to centrum, and

hypurals 3 + 4 fused and free from centrum; 54-1 no uroneural; 55-1 no stegural; 96-3 13 principal caudal-fin

rays; 97-2,3 two (in lower lobe) or three (in upper lobe) procurrent caudal-fin rays.
FIGURE 24.—Parazenidae

. Skeleton of Parazen pacificus, CAS 38404, 96 mm SL, lateral view. Character-state

(for selected unlabeled features not indicated in detailed osteological views for species of the family): 8-0 deep

lachrymal; 10-0 slender, tubular infraorbitals; 15-1 ascending premaxillary process reaching about one-third into

orbit; 44-0 neural spines oriented posterodorsally; 45-2 large vacuities in abdominal haemal spines; 46-0 abdomi-

nal haemal spines with prominent ventral processes; 47-1 ossified ribs only on last few abdominal vertebrae; 48-0

ossified epineurals on most abdominal vertebrae; 59-2 three dorsal-fin pterygiophores anterior to fourth neural

spine; 61-0 ventral end of shaft of first dorsal-fin pterygiophore in middle of first intemeural space, not contacting

first neural arch and spine; 63-1 symmetrical pterygiophores; 64-0 no supraneural: 73-1 dorsal-, anal-, and pecto-

ral-fin rays unbranched; 78-0 ventral end of supracleithrum simple; 81-0 pelvic fin about midway between anus

and pectoral fin; 99-1 one vacant intemeural space.

C>z85Ho
NUMBHR 618 67

80-0

FIGURF. 25— Parazenidae. Rear half of the skull and first vertebra of Parazen pacificus. USNM 187892, 110 mm

SL, lateral (left) and dorsal views. In the lateral view, the dermosphenotic and infraorbitals are removed, and the

upper regions of the suspensory shafts of the three pharyngobranchials are shown in their place of articulation to

the prootic (first pharyngobranchial) and to the cartilaginous area between the anterior end of the basioccipital,

posterior end of the parasphenoid, and lower border of the prootic (second and third pharyngobranchials). The

ossified epineural present on the first vertebra is not shown here or in Figure 26. Character-state: 1-0 parietal

present; 2-0 basisphenoid a long shaft; 6-0 supraocular serrations on frontal; 40-1 first neural arch and spine

closely applied to skull; 42-0 no long dorsal extension of first neural spine above attachment to skull; 77-0 no

supracleithral serrations; 78-0 ventral end of supracleithrum simple; 80-0 one long extrascapular.

3-0

FIGURE 26.—Parazenidae. Skull and first vertebra of Parazen pacificus, USNM 187892, 110 mm SL, ventral

view. The lateral ethmoids and nasals are removed. Character-state: 3-0 vomerine teeth present; 4-0 no parasphenoid opening into posterior myodome; 6-0 supraocular serrations on frontal.
68 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

3-0

FIGURE 27.—Parazenidae. Suspensorium and associated cranial bones and preopercle of Parazen pacificus, USNM 187892, 110 mm SL, lateral view. Character-state: 2-0 basisphenoid a long shaft; 3-0

vomerine teeth present; 18-1 palatine articulation pivotal; 19-1 reduced metapterygoid; 20-0 no symplectic

ventral flange.

FIGURE 28.—Parazenidae. Upper

(above) and lower jaws of Parazen

pacificus. USNM 187892, 110 mm

SL, lateral views. At the front of the

dentary, parallel lines show the basal

regions of attachment of the two dentary cartilages, which are removed; the

wavy lines and small circles in

mechanical stippling show cartilages

that attach to the inner surface of the

maxilla. The bands of teeth on the

inner surface along the anterior border

of the premaxilla cannot be seen in lateral view. Character-state: 13-0 postmaxillary process present; 14-0 alveolar process of premaxilla simple; 22-0

no dentary serrations (edge may be

irregular but not serrate).

22-0
NUMBER 618 69

37-1

33-1

FIGURE 29.—Parazenidae. Hyoid arch, branchiostegal rays, and, illustrated lowered from its natural position,

urohyal of Parazen pacificus, AMNH 29459, 105 mm SL, lateral view. Character-state: 33-1 beryciform foramen represented by deep groove on ceratohyal; 34-0 branchiostegals spread over/along both epihyal and ceratohyal; 35-0 ceratohyal notches present; 36-0 ceratohyal-epihyal articulation through cartilage only; 37-1 epihyal

less deep than rear of ceratohyal; 38-0 urohyal no longer than ceratohyal.
70 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

28-2

29-2

26-0

FIGURE 30.—Parazenidae. Right side of

the branchial arches (left), dorsal view,

artificially laid out with the dorsal elements reflected, and detail (upper right) of

the epibranchials and pharyngobranchials, artificially even more laid out,

reflected, and spread apart, of Parazen

pacificus, AMNH 29459, 105 mm SL.

Character-state: 26-0 no uncinate process;

27-0 no interarcual cartilage; 28-2 long

second pharyngobranchial suspensory

shaft; 29-2 long third pharyngobranchial

suspensory shaft; 30-0 first basibranchial

not depressed below surface of basihyal;

31-1 no fourth pharyngobranchial toothplate; 32-0 fifth ceratobranchial toothed.

FIGURE 31.—Parazenidae. Right cleithrum and the two postcleithra of Parazen

pacificus, USNM 187892, 110 mm SL,

medial view. Character-state: 75-0 postcleithrum in two pieces.
NUMBL:R 71

82-1

FIGURE 32.—Parazenidae. Basal region of the right pelvic

fin (left), ventral view, and pelvis, dorsal view, of Parazen

pacificus, USNM 187892, 110 mm SL. Character-state:

82-1 no pelvic-fin spine; 83-2 anterolateral processes as

broad flanges along basal region of rays; 84-0 no serrations on rays; 85-0 broadly overlapping medial processes

of basipterygia; 86-0 short posterior process of pelvis;

102-2 total of seven pelvic-fin elements.

67-0

FIGURE 33.—Parazenidae. Lower region of the first three anal-fin pterygiophores, single anal-fin spine, and first three anal-fin rays (left) and the upper

region of the first four dorsal-fin pterygiophores and first four dorsal-fin

spines of Parazen pacificus. CAS 38404, 96 mm SL, lateral views. The anteriorly protruding structure at the base of the second and third anal-fin rays is

the paired distal pterygiophore (a tiny pair of distal pterygiophores is also

between the two halves of the base of the first anal-fin ray but does not protrude). Character-state: 57-1 locking mechanism between first and second

dorsal-fin spines; 62-1 no ossified spinous dorsal-fin distal radial; 64-0 no

supraneural; 67-0 no fusion of first anal-fin spine to pterygiophore; 73-1

anal-fin rays unbranched; 100-1 one anal-fin spine. Unnumbered here and in

all of the other illustrations showing dorsal-fin spines is a synapomorphy of

all zeiforms: a single supernumerary spine on the first dorsal-fin pterygiophore.
72

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

51-1

49-0

54-1

55-1

53-1

97-7

FIGURE 34.—Parazenidae. Caudal skeleton of Parazen pacificus, USNM 187892, 110 mm SL, lateral view.

Character-state: 39-0 PU2 first modified vertebra in caudal peduncle; 49-0 NPU2 long; 50-0 hypurapophysis

present; 51-1 two epurals; 52-1 anterior end of parhypural slightly removed from and not embracing centrum;

53-1 hypurals 1 + 2 fused and fused to centrum, and hypurals 3+ 4 fused and free from centrum; 54-1 no uroneural; 55-1 no stegural; 96-3 13 principal caudal-fin rays; 97-7 seven procurrent caudal-fin rays both above and

below (polymorphic; about as often eight procurrents, 97-8).
NUMBER 618 73

63-1

FIGURE 35.-—Parazenidae. Lower region of the first

three anal-fin basal pterygiophores and the anal-fin

spine and first two anal-fin rays of Cyttopsis roseus,

AMNH 29460, 74 mm SL, lateral view. The anal-fin

spine is fused to the first pterygiophore in this specimen, and the first anal-fin ray, also borne on the first

pterygiophore, loses its segmentation in large specimens and becomes spine-like. Character-state: 63-1

symmetrical pterygiophores; 67-1 fusion of first analfin spine to pterygiophore only in some specimens (as

here); 73-1 anal-fin rays unbranched.

67-1

49-0

51-1

FIGURE 36.—Parazenidae. Caudal skeleton of Cyttopsis roseus, USNM 187873, 72 mm SL, lateral view.

Character-state: 39-1 PU3 first modified vertebra in

caudal peduncle; 49-0 NPU2 long; 50-1 no hypurapophysis; 51-1 two epurals; 52-1 anterior end of parhypural slightly removed from and not embracing centrum; 53-2 hypurals 1+2 + 3 + 4 fused and fused to

centrum; 54-1 no uroneural; 55-1 no stegural.

39-1
74

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 37.—Parazenidae. Right lachrymal, infraorbitals, and (detached from skull) dermosphenotic ofStefhopristes eos, USNM 226570, 91 mm SL, lateral view. Character-state: 5-1 honeycomb bone sculpturing; 8-0 deep

lachrymal; 9-1 three infraorbitals; 10-0 slender, tubular infraorbitals; 11-0 dermosphenotic a distinctly separate

ossification.

85-0

86-0

85-2

86-1 —

FIGURE 38.—Parazenidae and other families. Pelvis, dorsal view: A, Capromimus abbreviates (Zeniontidae),

AMS I.2365OO01, 61 mm SL; B, Stethopristes eos (Parazenidae), USNM 226570, 91 mm SL; C, Zenopsis conchifer (Zeidae), USNM 117280, 51 mm SL. Character-state: 85-0 broadly overlapping medial processes of

basipterygia; 85-2 basipterygia not in close contact in middle region; 86-0 short posterior process of pelvis; 86-1

long, rod-like posterior process of pelvis.
FIGURE 39.—Zeniontidae. Skeleton of Zenion hololepis, CAS 38409, 70 mm SL, lateral view. The pectoral fin

has been removed so as not to obscure underlying features. Character-state (for selected unlabeled features not

indicated in detailed osteological views for species of the family): 15-0 ascending premaxillary process reaching

no further than front of lateral ethmoid and orbit; 44-0 all neural spines oriented posterodorsally; 45-0 no vacuities in abdominal haemal spines; 46-0 abdominal haemal spines with prominent ventral processes; 47-1 ossified

ribs only on last abdominal vertebra; 48-0 ossified epineurals on most abdominal vertebrae; 58-1 two groups of

vacant intemeural spaces; 59-2 three dorsal-fin pterygiophores anterior to fourth neural spine; 63-1 symmetrical

pterygiophores; 73-1 dorsal- and anal-fin rays unbranched (also pectoral-fin rays, not shown); 78-0 ventral end

of supracleithrum simple; 81-1 pelvic fin slightly behind pectoral fin; 99-2 two vacant intemeural spaces.
76

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

6-0

FIGURE 40.—Zeniontidae. Rear half of the skull, first three vertebrae, supraneural, and first two dorsal-fin pterygiophores of Zenion hololepis, CAS 38409, 70 mm SL, lateral view. The dermosphenotic is removed, and the

ossified epineurals on the first and more posterior vertebrae are not shown. Character-state: 1-0 parietal present;

2-0 basisphenoid a long shaft; 4-0 no parasphenoid opening into posterior myodome; 6-0 supraocular serrations

on frontal; 40-1 first neural arch and spine closely applied to skull; 41-1 flexible articulation of second and next

few anterior abdominal vertebrae with skull-first vertebra, these vertebrae linked laterally and ventrally by ligamentous bands; 42-1 long dorsal extension of first neural spine above attachment to skull; 60-1 ventral end of

shaft of first dorsal-fin pterygiophore in first interneural/preneural space; 61-1 ventral end of shaft of first dorsalfin pterygiophore at front of first interneural/preneural space, contacting skull and first vertebra between the two

sides of first neural arch and spine; 64-1 one supraneural; 65-1 no cartilage at distal end of supraneural; 80-0 one

long extrascapular.
NUMBER 618 77

FIGURE 41.—Zeniontidae. Skull, dorsal (left) and ventral views, first two vertebrae, dorsal view, and first three

vertebrae, ventral view, of Zenion hololepis. CAS 38409, 70 mm SL. The supraneural is removed, and the ossified epineurals on the first and more posterior vertebrae are not shown. In the dorsal view, the parallel lines

between the first two vertebrae show where the ventral end of the shaft of the first dorsal-fin pterygiophore

(which is removed) is articulated; the parallel lines medially behind the ethmoid and between the frontals are cartilage. In the ventral view, parallel lines between the frontals and the pterosphenoids, and between the basisphenoid, exoccipitals, and prootics, are cartilage. Character-state: 1-0 parietal present; 3-0 vomerine teeth present

(teeth tiny; row of foramina lateral to each patch of teeth); 4-0 no parasphenoid opening into posterior myodome;

6-0 supraocular serrations on frontal; 11-0 dermosphenotic a distinctly separate ossification; 40-1 first neural

arch and spine closely applied to skull; 80-0 one long extrascapular.
78

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

22-2

FIGURE 42.—Zeniontidae. Lower jaw and associated cartilages of Zenion hololepis, USNM 307305, 48 mm SL,

lateral view. Parallel lines in front of the dentary show the proximal half of the more anterior of the two dentary

cartilages, rotated forward, and just behind it the proximal region of attachment of the more posterior dentary

cartilage, which has been removed (like the anterior cartilage, the posterior cartilage has a divided base). The

wavy lines and small circles in mechanical stippling at the top of the dentary and articular show cartilages and

ligaments that attach through a broad band of connective tissue along much of the posterior border of the maxilla.

Character-state: 21-2 two dental cartilages of moderate length, about equal or the first slightly shorter; 22-2 multiple dentary serrations.

20-1

FIGURE 43.—Zeniontidae. Suspensorium and preopercle of Zenion hololepis, CAS 38409, 70 mm SL, lateral

view, somewhat flattened out from its slightly oblique natural position to better expose the metapterygoid. Parallel lines at the posterior border of the palatine show its cartilaginous core. The extent of the ventral flange of the

symplectic below the upper border of the preopercle is indicated by a dashed line. Character-state: 18-1 palatine

articulation pivotal; 19-1 reduced metapterygoid; 20-1 symplectic ventral flange.
NUMBER 618 79

FIGURE 44.—Zeniontidae. Lachrymal and infraorbitals (excluding dermosphenotic) of Zenion hololepis, CAS

76856, 73 mm SL, lateral view. Specimens sometimes have one less infraorbital than illustrated, and the sixth

and seventh are sometimes consolidated. Character-state: 8-0 deep lachrymal; 9-2 six infraorbitals; 10-2 deep

infraorbitals with serrate vertical supporting flanges.

36-0

35-0

37-0

38-1

FIGURE 45.—Zeniontidae. Hyoid arch, branchiostegal rays, and urohyal of Zenion hololepis, CAS 38409, 70

mm SL, lateral view. Because the branchial apparatus is excessively flared in preserved specimens, the urohyal

is illustrated lower than its natural position. Character-state: 33-1 beryciform foramen represented by deep

groove on ceratohyal (ontogenetically polymorphic; beryciform foramen present at smaller sizes, 33-0); 34-0

branchiostegals spread over/along both epihyal and ceratohyal; 35-0 ceratohyal notches present; 36-0 ceratohyal-epihyal articulation through cartilage only; 37-0 epihyal about same depth as rear of ceratohyal; 38-1 urohyal large, longer than ceratohyal.
33-0

33-1

FIGURE 46.—Zeniontidae. Ontogenetic reduction of the

beryciform foramen in the ceratohyal of Zenion hololepis,

lateral views. It is represented in large specimens by a

deep groove along the surface of the bone: from top to

bottom, USNM 187864, 50 mm SL; USNM 307305, 48

mm SL; USNM 187864, 65 mm SL; USNM 187864, 73

mm SL. Character-state; 33-0 beryciform foramen

present; 33-1 beryciform foramen represented by deep

groove on ceratohyal.

FIGURE 47.—Zeniontidae. Dorsal view of the right side of the branchial arches (left), artificially laid out with the

dorsal elements reflected, and detail (upper right) of the epibranchials and pharyngobranchials, artificially even

more laid out, reflected, and spread apart, of Zenion hololepis. CAS 38409, 70 mm SL. Character-state: 26-0 no

uncinate process; 27-0 no interarcual cartilage; 28-2 long second pharyngobranchial suspensory shaft; 29-1 moderately long third pharyngobranchial suspensory shaft; 30-0 first basibranchial not depressed below surface of

basihyal; 31-1 no fourth pharyngobranchial toothplate; 32-0 fifth ceratobranchial toothed.
NUMBER 618 81

FIGURE 48.—Zeniontidae. Zenion hololepis, USNM 187864, 50 mm SL, lateral views: A, first three dorsal-fin

spines and pterygiophores; B, the two anal-fin spines and first anal-fin pterygiophore. Character-state: 57-3 locking mechanism between first, second, and third dorsal-fin spines; 62-1 no ossified spinous dorsal-fin distal

radial; 66-1 locking mechanism between first and second anal-fin spines; 68-1 second anal-fin spine less than

one-half length of first spine; 100-2 two anal-fin spines.
82

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 49.—Zeniontidae. Pelvis of Zenion hololepis, USNM

187864, 65 mm SL, dorsal view. The lateral extent of the overlapping medial flanges is indicated by dashed lines. Character-state:

85-0 broadly overlapping medial processes of basipterygia; 86-0

short posterior process of pelvis.

83-0

84-0

102-2

FIGURE 50.—Zeniontidae. Basal region of the right pelvic

fin of Zenion hololepis, USNM 187864, 73 mm SL, ventral view. See Figure 51 for details of the locking mechanism of the pelvic-fin spine against the pelvis. Characterstate: 82-0 pelvic-fin spine present; 83-0 no anterolateral

processes from bases of rays; 84-0 no serrations on rays;

102-2 total of seven pelvic-fin elements. m\

82-0
NUMBER618 83

FIGURE 51.—Zeniontidae. Zenion hololepis, USNM 187864, 50 mm SL, lateral (A) and posterior (slightly

oblique) (B) views: A, locking articulation between the roughened surface of the pelvic-fin spine and the pelvis;

B, detail of the deep indentation in the base of the pelvic-fin spine that rotates around a strong flange from the

anterior region of the articular facet on the pelvis. Small circles in mechanical stippling on the side of the pelvis

behind the pelvic-fin spine show the band of cartilage against which the pelvic-fin rays articulate. Characterstates are as in Figure 50.

FIGURE 52.—Zeniontidae. Right cleithrum and adjacent region of the single postcleithrum of Zenion

hololepis, CAS 38409, 70 mm SL, medial view. Character-state: 75-1 single postcleithrum; 76-0 no flange

on single postcleithrum.

76-0

W-75-1
84

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

97-3

51-1

\

49-0

96-3

FIGURE 53.—Zeniontidae. Caudal skeleton of Zenion hololepis, USNM 307305, 48 mm SL, lateral view. Character-state: 39-1 PU3 first modified vertebra in caudal peduncle; 49-0 NPU2 long; 50-1 no hypurapophysis; 51-1

two epurals; 52-1 anterior end of parhypural slightly removed from and not embracing centrum; 53-2 hypurals

1+2 + 3+ 4 fused and fused to centrum; 54-1 no uroneural; 55-1 no stegural; 96-3 13 principal caudal-fin rays;

97-2,3 two (in lower lobe) or three (in upper lobe) procurrent caudal-fin rays.
NUMBER 61S

85

FIGURE 54.—Zeniontidae. Jaws, suspensorium, and opercular bones (above) and lachrymal and infraorbitals

(excluding deimosphenotic) of Capromimus abbreviatus, AMS 1.23650001, 61 mm SL, lateral views. Characterstate: 8-0 deep lachrymal; 9-1 four infraorbitals; 10-0 slender, tubular infraorbitals; 13-0 postmaxillary process

present; 14-1 alveolar process of premaxilla ventrally indented into two blunt lobes; 18-1 palatine articulation

pivotal; 19-1 reduced metapterygoid; 20-0 no symplectic ventral flange; 22-2 multiple dentary serrations.
86

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 55.—Zeniontidae. Dentary of

Capromimus abbreviatus, AMS

1.23650001, 61 mm SL, lateral view.

Parallel lines and wavy lines show,

respectively, the more anterior and

more posterior dental cartilages in their

natural position. The distal end of the

posterior dental cartilage attaches to the

inner surface of the lower end of the

maxilla. Small circles in mechanical

stippling show the lower region of the

ligament from the upper ends of the

dentary and articular that attaches to the

inner surface of the middle of the maxilla. Character-state: 21-1 two dental

cartilages of moderate length, the first

shorter; 22-2 multiple dentary serrations.

21-1

28-2

29-1

FIGURE 56.—Zeniontidae. Right side of the branchial arches (left), artificially laid out with the dorsal elements

reflected, and detail (upper right) of the epibranchials and pharyngobranchials, artificially even more laid out,

reflected, and spread apart, of Capromimus abbreviatus, AMS 1.23650001, 61 mm SL, dorsal view. Characterstate: 26-0 no uncinate process; 27-0 no interarcual cartilage; 28-2 long second pharyngobranchial suspensory

shaft; 29-1 moderately long third pharyngobranchial suspensory shaft; 30-0 first basibranchial not depressed

below surface of basihyal; 31-1 no fourth pharyngobranchial toothplate; 32-0 fifth ceratobranchial toothed.
NUMBHR 618 87

FIGURE 57.—Zeniontidae. First four dorsal-fin

spines and first four dorsal-fin pterygiophores,

with an inset (top left) of the median flange on the

distal ends of the first two pterygiophores around

which the second dorsal-fin spine articulates (base

of second dorsal spine outlined by dashed line), of

Capromimus abbreviatus, AMS I.2365OOO1, 61

mm SL, lateral view. Character-state: 57-2 locking

mechanism between second and third dorsal-fin

spines; 62-1 no ossified spinous dorsal-fin distal

radial.

FIGURE 58.—Zeniontidae. The two anal-fin spines, first anal-fin

ray, and lower region of the first two anal-fin pterygiophores of

Capromimus abbreviatus, AMS 1.23650001, 61 mm SL, lateral

view. At the base of the anal-fin ray, the anteriorly protruding

structure is the paired distal pterygiophore. The region of the second anal-fin spine overlapped by the locking flange of the first

anal-fin spine is indicated by a dashed line. Character-state: 66-1

locking mechanism between first and second anal-fin spines: 68-1

second anal-fin spine less than one-half length of first spine; 100-

2 two anal-fin spines.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

49-0

51-2

FIGURE 59.—Zeniontidae. Caudal skeleton, lateral view: A, Cyttomimus stelgis, USNM 306156, 76 mm SL; B, Capromimus abbreviatus, AMS

1.23650001, 61 mm SL. Character-state: 39-1 PU3 first modified vertebra in

caudal peduncle (A, B); 49-0 NPU2 long (A, B); 50-0 hypurapophysis present;

50-1 no hypurapophysis; 51-2 one epural (A, B); 52-1 anterior end of parhypural slightly removed from and not embracing centrum (A, B); 53-2 hypurals

1+2 + 3 + 4 fused and fused to centrum (A, B); 54-1 no uroneural (A, B); 55-1

no stegural (A, B).
FIGURE 60.—Grammicolepididae.

Skeleton of Xenolepidichthys dalgleixhi, CAS 38403, 70 mm SL, lateral view. Character-state (for

selected unlabcled features not indicated in detailed osteological views

for species of the family): 8-1 moderately sized lachrymal; 10-0 slender, tubular infraorbitals; 42-1 long

dorsal extension of first neural spine

above attachment to skull; 44-1

some neural spines oriented anterodorsally; 45-0 no vacuities in

abdominal haemal spines; 46-0

abdominal haemal spines with

prominent ventral processes; 47-1

ossified rib only on ninth (penultimate) abdominal vertebra; 48-0

ossified epineurals on most abdominal vertebrae; 57-2 locking mechanism between second and third dorsal-fin spines; 58-1 two groups of

vacant interneural spaces; 61-1 ventral end of shaft of first dorsal-fin

pterygiophore at front of first interneural/preneural space, contacting

skull and first vertebra between the

two sides of first neural arch and

spine; 63-1 symmetrical pterygiophores; 64-1 one supraneural; 65-0

cartilage at distal end of supraneural;

66-1 locking mechanism between

first and second anal-fin spines; 73-1

dorsal, anal-, and pectoral-fin rays

unbranched; 78-0 ventral end of

supracleithrum simple; 81-2 pelvic

fin under pectoral fin; 99-4 four

vacant interneural spaces.
90

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

3-1

4-0

1-0

FIGURE 61.—Grammicolepididae. Skull and first vertebra of Xenolepidichthys dalgleishi, CAS 38406, 75 mm

SL, dorsal (left) and ventral views. Character-state: 1-0 parietal present; 3-1 no vomerine teeth; 4-0 no parasphenoid opening into posterior myodome; 6-0 supraocular serrations on frontal; 11-0 dermosphenotic a distinctly

separate ossification; 40-1 first neural arch and spine closely applied to skull; 80-0 one long extrascapular.
NUMBER 618 91

12-1

14-0

13-1

22-0

FIGURE 62.—Grammicolepididae. Jaws and cartilages in the ethmoid region of Xenolepidichthys dalgleishi,

USNM 320016, 64 mm SL, lateral view. Parallel lines show the cartilages below the rear end of the ascending

premaxillary process, between the lateral ethmoid and the vomer and parasphenoid, and along the upper rear border of the articular. Wavy lines show the cartilage that continues posteriorly to the rear of the orbit and contacts

the pterosphenoids, a probable synapomorphy of zeiforms not enumerated herein because the condition is

unknown in many taxa (for our documentation of this, see character VI of Johnson and Patterson, 1993, in "Previous Phylogenetic Analyses of Zeiforms")- Character state: 3-1 no vomerine teeth; 12-1 small mouth, alveolar

process less deep than lateral ethmoid; 13-1 no postmaxillary process; 14-0 alveolar process of premaxilla simple; 15-0 ascending premaxillary process reaching no further than front of lateral ethmoid and orbit; 22-0 no dentary serrations.
92

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

20-1

FIGURE 63.—Grammicolepididae. Suspensorium and preopercle of Xenolepidichthys dalgleishi, CAS 38406, 75

mm SL, lateral view. The extent of the ventral flange of the symplectic below the upper border of the preopercle

is indicated by a dashed line, as is its anterior end under the quadrate. Character-state: 18-1 palatine articulation

pivotal; 19-1 reduced metapterygoid; 20-1 symplectic ventral flange.
NUMBER 618 93

FIGURE 64.—Grammicolepididae. Lower jaw of Xenolepidichthys dalgleishi, USNM 320016, 59 mm SL, lateral view. The

illustration is at a right angle to the lower jaw, whereas the view

of the lower jaw in Figure 62 is somewhat oblique because of

the considerable inward slant of the lower border in its natural

position. Small circles in mechanical stippling show cartilages

along the upper border of the articular. Character-state: 22-0 no

dentary serrations.

FIGURE 65.—Grammicolepididae. Hyoid arch, branchiostegal rays, and urohyal of Xenolepidichthys dalgleishi,

CAS 38406, 75 mm SL, lateral view. Character-state: 33-0 beryciform foramen present; 34-0 branchiostegals

spread over/along both epihyal and ceratohyal; 35-1 no ceratohyal notches; 36-0 ceratohyal-epihyal articulation

through cartilage only; 37-1 epihyal less deep than rear of ceratohyal; 38-0 urohyal no longer than ceratohyal.
94 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

29-0 31-1

30-1

FIGURE 66.—Grammicolepididae. Xenolepidichthys dalgleishi, CAS 38406, 75 mm SL, dorsal (A, B) and lateral

(C) views: A, right side of the branchial arches, artificially laid out with the dorsal elements reflected; B, detail of

the epibranchials and pharyngobranchials, artificially even more laid out, reflected, and spread apart; C, basihyal

(right) and the three basibranchials (first to third, from front (below) to rear (above)). (Scale bar the same for A

and C.) Character-state: 26-0 no uncinate process; 27-0 no interarcual cartilage; 28-0 short second pharyngobranchial suspensory shaft; 29-0 short third pharyngobranchial suspensory shaft; 30-1 first basibranchial depressed

below surface of basihyal; 31-1 no fourth pharyngobranchial toothplate; 32-1 no fifth ceratobranchial teeth.
NUMBF.R618 95

FIGURE 67.—Grammicolepididae. Xenolepidichihys

dalgleishi, CAS 38406, 75 mm SL: A, right cleithrum

and adjacent region of the single postcleithrum, medial

view; B, pelvis, dorsal view. Character-state: 75-1 single

postcleithrum; 76-0 no flange on single postcleithrum;

85-2 basipterygia not in close contact in middle region;

86-0 short posterior process of pelvis.

FIGURE 68.—Grammicolepididae. Basal region of the right

pelvic fin of Xenolepidichihys dalgleishi. CAS 38406, 75

mm SL, ventral view. Character-state: 82-0 pelvic-fin

spine present; 83-0 no anterolateral processes from bases

of rays; 84-1 serrations present on low crests along posteroventral edges of rays; 102-2 total of seven pelvic-fin elements.
96

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

97-1

51-1

49-0 96-1

50-0

53-2

FIGURE 69.—Grammicolepididae. Caudal skeleton of Xenolepidichlhys dalgleishi, CAS 38406, 75 mm SL, lateral view. Character-state: 39-1 PU3 first modified vertebra in caudal peduncle; 49-0 NPU2 long; 50-0

hypurapophysis present (relatively low and short); 51-1 two epurals; 52-2 peg and socket joint; 53-2 hypurals 1 +

2 + 3+ 4 fused and fused to centrum; 54-1 no uroneural; 55-1 no stegural; 96-1 15 principal caudal-fin rays; 97-

I one procurrent caudal-tin ray, both above and below.
FIGURE 70.—Grammicolepididae. Skeleton of Macrurocyttus acanthopodus, MNHN 1994-1073,45 mm SL, lateral view, based on a radiograph. The neural spine of the ninth caudal vertebra is abnormally bifurcate. Character-state (for selected unlabeled features not indicated in detailed osteological views for this species): 15-0

ascending premaxillary process reaching no furher than front of lateral ethmoid and orbit; 44-1 some neural

spines oriented anterodorsally; 45-0 no vacuities in abdominal haemal spines; 46-1 abdominal haemal spines

without prominent ventral processes; 47-1 ossified rib only on thirteenth (last) abdominal vertebra (minority condition, in two of seven specimens; usually no ossified ribs, 47-3); 48-3 no ossified epineurals; 58-1 two groups of

vacant interneural spaces (minority condition; usually three groups, 58-2); 59-2 three dorsal-fin pterygiophores

anterior to fourth neural spine; 61-1 ventral end of shaft of first dorsal-fin pterygiophore at front of first intemeural/preneural space, contacting skull and first vertebra between the two sides of first neural arch and spine; 63-1

symmetrical pterygiophores; 64-1 one supraneural; 81-2 pelvic fin somewhat anterior to pectoral fin; 99-5 five

vacant interneural spaces; 100-0 no anal-fin spine.
98

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

4-0

42-1

\

4-0

FIGURE 71.—Grammicolepididae. Skull and first vertebra of Macrurocyttus acanthopodus, USNM 93144, disarticulated specimen of unknown SL (see "Material"), dorsal (A), ventral (B), and lateral (c) views. Small circles in

mechanical stippling show some of the major cartilages, with open spaces between the bones of the rear of the

skull also cartilaginous. Character-state: 1-1 no parietal; 2-2 no basisphenoid; 3-1 no vomerine teeth; 4-0 no

parasphenoid opening into posterior myodome; 6-0 supraocular serrations on frontal; 40-1 first neural arch and

spine closely applied to skull; 42-1 long dorsal extension of first neural spine above attachment to skull.
NUMRF.R6I8 99

36-0

B

37-0

103-2

97-0

54-1

55-1

53-5

FIGURE 72.—Grammicolepididae. Macrurocyttus acanthopodus: A, USNM 93144, pelvis of disarticulated

specimen of unknown SL (see "Material"), dorsal view. B, c, MNHN 1994-1072, 43 mm SL, lateral view: B,

hyoid arch and branchiostegals; c, caudal skeleton. Character-state: 33-1 beryciform foramen represented by

deep groove on ceratohyal; 34-0 branchiostegals spread over/along both epihyal and ceratohyal; 35-1 no ceratohyal notches; 36-0 ceratohyal-epihyal articulation through cartilage only; 37-0 epihyal about same depth as rear

of ceratohyal; 39-0 PU2 first modified vertebra in caudal peduncle; 49-0 NPU2 long; 50-1 no hypurapophysis;

51-2 one epural; 52-1 anterior end of parhypural slightly removed from and not embracing centrum; 53-5 all

hypurals fused to centrum (fifth hypural indistinguishably fused to the others or absent, in either case a condition unique among zeiforms, although fifth hypural with partial fusion in some zeids); 54-1 no uroneural; 55-1

no stegural; 85-0 broadly overlapping medial processes of basipterygia; 86-0 moderately long posterior process

of pelvis; 96-3 13 principal caudal-fin rays; 97-0 no procurrent caudal-fin rays above or below; 103-2 six branchiostegals (2+4). There is only one hypohyal, a condition unique among zeiforms, but the foramen in the

largely cartilaginous element may indicate a compound origin of the single piece.
FIGURE 73.—Zeidae. Skeleton ofZeusfaber, USNM 320063, 80

mm SL, lateral view. Character-state (for selected unlabeled features not

indicated in detailed osteological views for species of the family): 8-1 moderately sized lachrymal; 10-0 slender, tubular infraorbitals; 15-0 ascending

premaxillary process reaching no further than front of lateral ethmoid and

orbit; 44-1 some neural spines oriented anterodorsally; 45-0 no vacuities in

abdominal haemal spines; 46-0 abdominal haemal spines with prominent

ventral processes; 48-1 ossified epineurals on only a few anterior abdominal vertebrae; 57-3 locking mechanism between first, second, and third

dorsal-fin spines; 58-2 three groups of vacant intemeural spaces (polymorphic; as often four groups, 58-3); 75-1 single postcleithrum; 78-1 ventral

end of supracleithrum deeply bifurcate; 81-2 pelvic fin anterior to pectoral

fin; 99-5 five vacant intemeural spaces.
NUMBER 618 101

60-1

61-1

62-1

80-1

FIGURE 74.—Zeidae. Rear of the skull and the first six vertebrae and the more anterior dorsal-fin pterygiophores

and dorsal-fin spines of Zenopsis conchifer, CAS 47401, 54 mm SL, lateral view. Parallel lines show the cartilaginous ventral ends of the pterygiophores and the bands of connective tissue between the anterior few vertebrae

(which appear as ventral straps in lateral view under transmitted light but are actually much broader bands).

Character-state: 1-0 parietal present; 4-0 no parasphenoid opening into posterior myodome; 11-1 dermosphenotic

consolidated with sphenotic; 40-1 first neural arch and spine closely applied to skull; 41-1 flexible articulation of

second and next few anterior abdominal vertebrae with skull-first vertebra, these vertebrae linked laterally and

ventrally by ligamentous bands; 47-0 ossified ribs on most abdominal vertebrae behind fourth; 48-0 ossified

epineurals on most abdominal vertebrae; 59-3 four dorsal-fin pterygiophores anterior to fourth neural spine; 60-1

ventral end of shaft of first dorsal-fin pterygiophore in first interneural/preneural space; 61-1 ventral end of shaft

of first dorsal-fin pterygiophore at front of first interneural/preneural space, contacting skull and first vertebra

between the two sides of first neural arch and spine; 62-1 no ossified spinous dorsal-fin distal radial; 64-0 no

supraneural; 80-1 two tubular extrascapulars, not closely held to skull.
6-0 —

i n

FIGURE 75.—Zeidae. Rear of the skull and first vertebra of Zenopsis

conchifer, CAS 47401, 54 mm SL, obliquely posterior view. Parallel

lines between the bifid neural arch and spine of the first vertebra

show the position of the base of the first dorsal-fin pterygiophore,

which has been removed. The dorsal ends of the cleithrum and supracleithrum are shown on the left side. Character-state: 1-0 parietal

present; 6-0 supraocular serrations on frontal (polymorphic; with

increasing size, no serrations, 6-1); 40-1 first neural arch and spine

closely applied to skull; 61-1 ventral end of shaft of first dorsal-fin

pterygiophore at front of first intemeural/preneural space, contacting

skull and first vertebra between the two sides of first neural arch and

spine; 80-1 two tubular extrascapulars, not closely held to skull.

FIGURE 76.—Zeidae. Skull of Zenopsis nebulosus, AMNH 92291, 325 mm SL, dorsal (left) and ventral

views. Character-state: 1-0 parietal present; 3-0 vomerine teeth present; 4-0 no parasphenoid opening into

posterior myodome; 6-0 supraocular serrations on frontal (many more serrations at smaller sizes, but

polymorphic; serrations lost at largest sizes, 6-1); 40-1 first neural arch and spine closely applied to skull;

80-1 two tubular extrascapulars, not closely held to skull.
NUMBER 618 103

FIGURE 77.—Zeidae. Left-side jaws of Zenopsis nebulosus,

AMNH 92291, 325 mm SL, medial view, and detail (left) of

the vomer, ventral view. Character-state: 3-0 vomerine teeth

present; 13-0 postmaxillary process present; 14-2 alveolar process of premaxilla deeply bifurcate ventrally; 22-0 no dentary

serrations.

20-0

FIGURE 78.—Zeidae. Right-side suspensorium of Zenopsis

conchifer, CAS 47401, 54 mm SL, lateral view. Characterstate: 18-1 palatine articulation pivotal; 19-1 reduced metapterygoid; 20-0 no symplectic ventral flange.
104 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

34-0

FIGURE 79.—Zeidae. Hyoid arch,

branchiostegal rays, basihyal, and

first two basibranchials of Zenopsis

conchifer, CAS 47401, 54 mm SL,

lateral view. The urohyal is illustrated lower and forward of its natural position internal to the hypohyals and anterior end of the

ceratohyal. Character-state: 33-0

beryciform foramen present; 34-0

branchiostegals spread over/along

both epihyal and ceratohyal; 35-0

ceratohyal notches present; 36-2

ceratohyal-epihyal articulation with

bony interdigitations at all sizes;

37-1 epihyal less deep than rear of

ceratohyal; 38-0 urohyal no longer

than ceratohyal.

28-2

29-1

27-0

31-1

26-0

FIGURE 80.—Zeidae. Right side of the branchial arches

(left), dorsal view, artificially laid out with the dorsal elements reflected, and detail (upper right) of the epibranchials and pharyngobranchials, artificially even more laid

out, reflected, and spread apart, of Zenopsis conchifer,

FMNH 67090, 44 mm SL. Character-state: 26-0 no uncinate process; 27-0 no interarcual cartilage; 28-2 long second pharyngobranchial suspensory shaft; 29-1 moderately

long third pharyngobranchial suspensory shaft; 30-0 first

basibranchial not depressed below surface of basihyal; 31-

I no fourth pharyngobranchial toothplate; 32-0 fifth ceratobranchial toothed.
NUMBER 618 105

73-1

63-

FlGURE 81.—Zeidae. Typical soft

dorsal-fin pterygiophores (between

neural spines of third and fourth

abdominal vertebrae) (left), and

detail of the articulation of the base

of a dorsal-fin ray with the distal

end of the pterygiophore in Zenopsis conchifer, FMNH 67090, 44

mm SL, lateral views. Parallel lines

are cartilages. The dashed line at

the base of the dorsal-fin ray in the

detail is the ossified distal pterygiophore. Character-state: 63-1 symmetrical pterygiophores; 73-1 dorsal-fin rays unbranched.

62-1

FIGURE 82.—Zeidae. First three spinous dorsal-fin pterygiophores (fourth incomplete), with the first two buckler

plates in place on the right side, of Zenopsis conchifer,

CAS 47401,54 mm SL, dorsal view. The first three dorsalfin spines articulate around the median crests on each

pterygiophore (toward middle of first pterygiophore and

front of second and third pterygiophores). Character-state:

62-1 no ossified spinous dorsal-fin distal radial; 88-3 large

buckler plates along base of anterior dorsal-fin spines.

66-1

FIGURE 83.—Zeidae.

The three anal-fin spines,

first anal-fin ray, and first

three pterygiophores of Zenopsis

conchifer, CAS 47401, 54 mm SL,

lateral view. Character-state: 66-1 locking mechanism between first and second

anal-fin spines; 68-0 second anal-fin spine more

than one-half length of first spine; 73-1 anal-fin rays

unbranched; 100-3 three anal-fin spines.
106

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

102-3

FIGURE 84.—Zeidae. Left-side pelvis and pelvic fin (anterior to right) of Zenopsis conchifer, CAS 47401, 54 mm

SL, medial view, with detail (right) of the ray-like first element of the pelvic-fin (lateral and medial halves separate throughout their lengths and unbranched, but unsegmented), anterior view. Character-state: 82-1 no pelvicfin spine; 86-1 long, rod-like posterior process of pelvis; 102-3 total of six pelvic-fin elements.
NUMBER 618 107

FIGURE 85.—Zeidae. Basal region of the right pelvic fin of Zenopsis conchifer, USNM 117280, 81 mm SL, lateral (A) and medial (B) views. Parallel lines show cartilage. (Anterior to right in A, to left in B.) Character-state:

82-1 no pelvic-fin spine; 83-1 anterolateral processes as prongs from bases of rays; 84-0 no serrations on rays;

102-3 total of six pelvic-fin elements.

97-1

51-2

39-0

49-0

96-3

FIGURE 86.—Zeidae. Caudal

skeleton of Zenopsis conchifer,

CAS 47401, 54 mm SL, lateral

view. Character-state: 39-0 PU2

first modified vertebra in caudal

peduncle; 49-0 NPU2 long; 50-1

no hypurapophysis; 51-2 one

epural; 52-1 anterior end of parhypural slightly removed from

and not embracing centrum; 53-2

hypurals 1 + 2 + 3+ 4 fused and

fused to centrum; 54-1 no uroneural; 55-1 no stegural; 56-1

extra-caudal ossicle present; 96-3

13 principal caudal-fin rays; 97-1

one procurrent caudal-fin ray

above and below.

53-2
108

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

56-1

"v _ _: ym

FIGURE 88.—Zeidae. Variation in the caudal skeleton of Zeus faber, lateral

views: bottom, USNM 320014, 59 mm SL (most common condition); top,

USNM 307842, 67 mm SL. Character-state: 56-1 extra-caudal ossicle present

(USNM 307842).

FIGURE 87.—Zeidae. Variation in the caudal skeleton of Zenopsis nebulosus,

lateral views: A, AMNH 92291, 325 mm SL (most common condition); B,

AMNH 95024, 335 mm SL; C, AMNH 95028, 340 mm SL. Character-state:

56-1 extra-caudal ossicle present (B, C).
FIGURE 89.—Fossil Zeiformes. Skeleton of Cretazeus rinaldi Tyler et al. (2001), holotype, Collezione della Citta

di Nardo, CCN 6659C, 53 mm SL, lateral view. This is the earliest known zeiform, from the Upper Cretaceous of

Nardo, Italy (about 72 m.y.a.), and, like the two morphologically primitive taxa of early zeiforms from the upper

Paleocene, it cannot be placed in any of the extant families. Cretazeus already exhibits such derived zeiform features as 18-1 palatine articulation pivotal; 19-1 reduced metaptcrygoid; 40-1 first neural arch and spine closely

applied to skull (better seen in other specimens); 44-1 some neural spines oriented anterodorsally; 49-0 NPU2

long; 52-1 anterior end of parhypural slightly removed from and not embracing centrum; 53-2 hypurals 1+2 + 3

+ 4 fused and fused to centrum; 54-1 no uroneural; 58-0, one group of vacant interneural spaces (variable

between the five type specimens, two vacancies in one group in this specimen); 60-1 first dorsal-fin pterygiophore attached to skull at first neural arch and spine; 63-1 symmetrical pterygiophores; 64-1 one supraneural; 73-

1 unbranched dorsal-, anal-, and pectoral-fin rays; 88-4 large buckler plates mid-abdominally from pelvic fin to

anus (independent of Hoplostethus); 11 principal caudal-fin rays, even fewer than the 13 of nearly all exant zeiforms (96-3). The total number of vertebrae in this species is 28 or (this specimen) 29; single supernumerary dorsal-fin spine on first ptcrygiophore (not enumerated).
10

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 90.—Fossil Zeiformes. Skeleton of Zenopsis clarus Danilchenko (1960), holotype, Paleontological Institute, Moscow, PIN 1413-22, 25 mm SL, lateral view. This is one of the several taxa of zeiforms from the Oligocene (in this case from the Caucasus, Russia), all of which clearly are members of the Zeidae and can be

assigned to either Zenopsis or Zeus (Baciu, Bannikov, and Tyler, in prep.). In addition to most of the characters

given in the legend for Figure 89 of the Upper Cretaceous Cretazeus (except supraneural absent, different buckler arrangement, 34 or 35 vertebrae, and 13 caudal-fin rays), the Oligocene taxa of Zenopsis have the following

zeid features: 14-2 alveolar process of premaxilla deeply bifurcate ventrally; 47-0 ossified ribs on most abdominal vertebrae behind fourth; 59-3 four dorsal-fin pterygiophores anterior to fourth neural spine; 79-1 cleithral

process present; 86-1 long, rod-like posterior process of pelvis; 88-3 large buckler plates along dorsal- and analfin bases and mid-abdominally; 94-5 14 abdominal vertebrae; 98-6 10 dorsal-fin spines; 102-3 total of six pelvicfin elements.


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