24 Gnathostome Fishes

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M. L. J. Stiassny

E. O. Wiley

G. D. Johnson

M. R. de Carvalho

410

Gnathostomata are a species-rich assemblage that, with the

exclusion of the Petromyzontiformes (lampreys, 45 spp.), represents

all living members of Vertebrata. Gnathostomes are

most notably characterized by the possession of endoskeletal

jaws primitively formed of dorsal palatoquadrate and ventral

Meckelian cartilages articulating at a mandibular joint. Our task

here is to provide a review of a large (paraphyletic) subset of

gnathostome diversity—an artificial grouping often referred

to as the “jawed fishes”: chondrichthyans, “piscine sarcopterygians,”

and actinopterygians. We treat all living jawed vertebrates

with the exclusion of most Sarcopterygii—the

tetrapods—since they are discussed in other chapters. After a

review of the chondrichthyans or cartilaginous fishes and a

brief summary of the so-called “piscine sarcopterygians,” we

focus our contribution on the largest and most diverse of the

three groups, the Actinopterygii, or rayfin fishes.

As a guide to the chapter, figure 24.1 presents, in broad

summary, our understanding of the interrelationships among

extant gnathostome lineages and indicates their past and

present numbers (with counts of nominal families indicated by

column width through time). Much of the stratigraphic information

for osteichthyans is from Patterson (1993, 1994), and

that for chondrichthyans is mostly from Cappetta et al. (1993).

Chondrichthyes (Cartilaginous Fishes)

Chondrichthyans (sharks, rays, and chimaeras) include approximately

1000 living species (Compagno 1999), several

dozen of which remain undescribed. Recent sharks and rays

are further united in the subclass Elasmobranchii (975+ spp.),

whereas the chimaeras form the subclass Holocephali (35+

spp.). All chimaeras are marine; as are most sharks and rays,

but about 15 living elasmobranch species are euryhaline, and

some 30 are permanently restricted to freshwater (Compagno

and Cook 1995).

Chondrichthyans are characterized by perichondral prismatic

calcification; the prisms form a honeycomb-like mosaic

that covers most of the cartilaginous endoskeleton

(Schaeffer 1981, Janvier 1996). Paired male intromittent

organs derived from pelvic radials (claspers) are probably

another chondrichthyan synapomorphy, although they are

unknown in some early fossil forms (e.g., the Devonian

Cladoselache and Carboniferous Caseodus), but all recent

chondrichthyans and most articulated fossil taxa have them

(Zangerl 1981). Earlier notions that sharks, rays, and chimaeras

evolved independently from placoderm ancestors (Stensiц

1925, Holmgren 1942, Шrvig 1960, 1962; Patterson 1965),

culminating in the Elasmobranchiomorphi (placoderms +

chondrichthyans) of Stensiц (e.g., 1958, 1963, 1969) and

Jarvik (e.g., 1960, 1977, 1980), have not survived close inspection

(e.g., Compagno 1973, Miles and Young 1977);

chondrichthyan monophyly is no longer seriously challenged

(Schaeffer 1981, Maisey 1984).

Sharks, rays, and chimaeras form an ancient lineage. The

earliest putative remains are dermal denticles from the Late

Ordovician of Colorado [some 450 million years ago (Mya)];

the first braincase is from the Early Devonian of South AfGnathostome

Fishes 411

rica some 60 million years later (Maisey and Anderson 2001).

The divergence between elasmobranchs and holocephalans

is also relatively old, because isolated holocephalan tooth

plates are known from the Late Devonian (Zangerl 1981,

Stahl 1999), and articulated specimens from the Early Carboniferous

(320 Mya; Lund 1990, Janvier 1996). A few of

the earliest known fossil sharks may be basal to the elasmobranch–

holocephalan dichotomy, such as Pucapampella from

the Devonian of Bolivia (Maisey 2001), but much work remains

to be done in early chondrichthyan phylogeny (Coates

and Sequeira 1998). Sharks were remarkably diverse morphologically

and ecologically during much of the Paleozoic,

considerably more so than early bony fishes. Some 32 families

existed during the Carboniferous, but many of these went

extinct before the end of the Permian (Cappetta et al. 1993;

fig. 24.1).

The entrenched notion that sharks are primitive or ancestral

vertebrates because of their antiquity, “generalized

design,” and lack of endochondral (cellular) bone (e.g.,

Dean 1895, Woodward 1898) is contradicted by the theory

that bone may have been lost in sharks, because it is widely

distributed among stem gnathostomes (Stensiц 1925,

Maisey 1986). Furthermore, acellular bone is present in the

dorsal spine-brush complex of an early shark (Stethacanthus;

Coates et al. 1999) and also in the teeth, denticles, and

vertebrae of extant chondrichthyans (Kemp and Westrin

1979, Hall 1982, Janvier 1996), supporting the assertion

that sharks evolved from bony ancestors. Highly complex,

derived attributes of elasmobranchs, such as their semicircular

canal arrangement (Schaeffer 1981), internal fertilization,

and formation of maternal–fetal connections

(“placentas” of some living forms; Hamlett and Koob 1999),

Figure 24.1. Current estimate

of relationships among extant

gnathostome lineages. Past and

present counts of nominal

families are indicated by column

width through time (tetrapod

diversity truncated, chondrichthyan

diversity truncated to the

left, and acanthomorph diversity

truncated to the right). Stratigraphic

information for

Osteichthyes is taken from

Patterson (1993, 1994) but with

new data for Polypteriformes

from Dutheil (1999) and for

Otophysi from Filleul and

Maisey (in press). Data for

Chondrichthyes are from

Cappatta et al. (1993), with

complementary information

from Janvier (1996) and other

sources. For practical reasons,

familial diversity is charted and

this does not necessarily reflect

known species diversity.

412 The Relationships of Animals: Deuterostomes

reveal, in fact, that sharks are much more “advanced” than

previously thought.

Elasmobranchs (Sharks and Rays)

Modern sharks and rays share with certain Mesozoic fossils

(e.g., Palaeospinax, Synechodus) calcified vertebrae and specialized

enameloid in their teeth (both secondarily lost in

some living forms) and are united with them in Neoselachii

(Schaeffer 1967, 1981, Schaeffer and Williams 1977, Maisey

1984). Most of modern elasmobranch diversity originated in

the Late Cretaceous to Early Tertiary (some 55–90 Mya), but

several extant lineages have fossil members, usually represented

by isolated teeth, dating back to the Early Jurassic

(some 200 Mya).

Recent phylogenetic studies have recognized two major

lineages of living elasmobranchs, Galeomorphi (galeomorph

sharks) and Squalomorphi (squalomorphs or squaleans;

Shirai 1992, 1996, Carvalho 1996; fig. 24.2). These studies,

however, differ in the composition of Hexanchiformes and

Squaliformes, and in relation to the coding and interpretation

of many features; the tree adopted here (fig. 24.2) is

modified from Carvalho (1996).

The phylogeny in figure 24.2 is the most supported by

morphological characters, but an alternative scheme has been

proposed on the basis of the nuclear RAG-1 gene ( J. G.

Maisey, pers. comm.), in which modern sharks are monophyletic

without the rays (an “all-shark” hypothesis). Stratigraphic

data are slightly at odds with both hypotheses, but

more so with the morphological one, because there are no

Early Jurassic squaloids, pristiophoroids, or squatinoids. But

lack of stratigraphic harmony will persist unless these taxa

are demonstrated to comprise a crown group within a monophyletic

“all-shark” collective (i.e., with galeomorphs basal

to them). Nonetheless, dozens of well-substantiated morphological

characters successively link various shark and

all batoid groups in Squalomorphi, many of which would

have to be overturned if sharks are to be considered monophyletic

to the exclusion of rays.

Historically, some of the difficulties in discerning relationships

among elasmobranchs have been due to the highly

derived design of certain taxa (e.g., angelsharks, sawsharks,

batoids, electric rays), which has led several workers (e.g.,

Regan 1906, Compagno 1973, 1977) to isolate them in their

own lineages, ignoring their homologous features shared with

other elasmobranch groups (Carvalho 1996). Elevated levels

of homoplasy (Shirai 1992, Carvalho 1996, McEachran

and Dunn 1998), coupled with the lack of dermal ossifications

(a plentiful source of systematically useful characters

in bony fishes), hinders the recovery of phylogenetic patterns

within elasmobranchs. Moreover, the (erroneous) notion that

there is nothing left to accomplish in chondrichthyan systematics

is unfortunately common. In fact, the situation is

quite the contrary, because many taxa are only “phenetically”

defined and require rigorous phylogenetic treatment (e.g.,

within Carcharhiniformes and Myliobatiformes). However,

many morphological complexes still require more in-depth

descriptive and comparative study (in the style of Miyake

1988, Miyake et al. 1992) before they can be confidently used

in phylogenetic analyses.

The general morphology, physiology, and reproduction

of extant sharks and rays are comprehensively reviewed in

Hamlett (1999). Fossil forms are discussed in Cappetta

(1987) and Janvier (1996). Below is a brief account of ex-

Figure 24.2. Intrarelationships

of extant chondrichthyan

lineages based mostly on

Carvalho (1996). Relationships

among rays (Batoidea) are left

unresolved, with guitarfishes

(Rhinobatiformes) in quotation

marks because the group is

probably not monophyletic (see

McEachran et al. 1996).

Gnathostome Fishes 413

tant elasmobranch orders; their monophyly ranges from the

relatively well established (Orectolobiformes) to the poorly

defined (Squaliformes; Compagno 1973, 1977, Shirai 1996,

Carvalho 1996).

Galeomorph sharks encompass four orders (fig. 24.2):

Heterodontiformes (bullhead sharks), Orectolobiformes

(carpet sharks), Lamniformes (mackerel sharks), and Carcharhiniformes

(ground sharks). Galeomorphs have various specializations

(Compagno 1973, 1977), such as the proximity

between the hyomandibular fossa and the orbit on the neurocranium,

and are the dominant sharks of shallow and epipelagic

waters worldwide (Compagno 1984b, 1988, 2001).

The two most basal galeomorph orders are primarily

benthic, inshore sharks. Bullheads (Heterodontus, eight spp.)

are distributed in tropical and warm-temperate seas of the

western and eastern Pacific Ocean and western Indian Ocean

(Compagno 2001). Heterodontus has a unique dentition,

composed of both clutching and grinding teeth, and is oviparous.

It was once believed to be closely related to more primitive

Mesozoic hybodont sharks (which also had dorsal fin

spines) and therefore regarded as a living relic (e.g., Woodward

1889, Smith 1942), but its ancestry with modern

(galeomorph) sharks is strongly corroborated (Maisey 1982).

Orectolobiforms (14 genera, 32+ spp.) are among the most

colorful elasmobranchs, occurring in tropical to warm-temperate

shallow waters; they are most diverse in the Indo-West

Pacific region but occur worldwide. Species are aplacentally

viviparous or oviparous. One orectolobiform, the planktophagous

whale shark (Rhincodon typus), is the largest known

fish species, reaching 15 m in length. Derived characters of

carpet sharks include their complete oronasal grooves and

arrangement of cranial muscles (Dingerkus 1986, Goto 2001).

Their taxonomy is reviewed in Compagno (2001), and their

intrarelationships in Dingerkus (1986) and Goto (2001). An

alternative view recognizes bullheads and carpet sharks as sister

groups (Compagno 1973; fig. 24.2).

From a systematic perspective, Lamniformes (10 genera,

15 spp.) contain some of the best-known sharks, characterized

by their “lamniform tooth pattern” (Compagno

1990, 2001). Although their low modern-day diversity pales

compared with the numerous Cretaceous and Tertiary species

described from isolated teeth (Cappetta 1987), this

order contains some of the most notorious sharks, such as

the great white (Klimley and Ainley 1997), its gigantic fossil

cousin Carcharodon megalodon (Gottfried et al. 1996), the

megamouth (now known from some 15 occurrences worldwide;

Yano et al. 1997), and the filter-feeding basking shark.

Lamniforms are yolk-sac viviparous, and adelphophagy

(embryos consuming each other in utero) and oophagy (embryos

eating uterine eggs) have been documented in some

species (Gilmore 1993). Molecular data sets (Naylor et al.

1997, Morrissey et al. 1997) are at odds with morphological

ones (and with each other), indicating that the jury is

still out in relation to the evolutionary history of lamniform

genera.

Carcharhiniformes (48 genera, 216+ spp.) are by far the

largest order of sharks, containing more than half of all living

species, and about half of all shark genera (Compagno

1984b). Carcharhiniforms have specialized secondary lower

eyelids (nictitating eyelids), as well as unique clasper skeletons

(Compagno 1988). Species are oviparous (Scyliorhinidae)

or viviparous, with or without the development of a

yolk-sac placenta (Hamlett and Koob 1999). Ground sharks

range from sluggish, bottom-dwelling catsharks (Scyliorhinidae,

the largest shark family) to epipelagic, streamlined,

and active requiem sharks (Carcharhinidae), which includes

some of the most common and economically important species

(e.g., blue and tiger sharks, Carcharhinus spp.). Hammerhead

sharks (Sphyrnidae) are morphologically very

distinctive (Nakaya 1995) and capable of complex behavioral

patterns (e.g., Myrberg and Gruber 1974). Some ground

sharks may be restricted to freshwater (Glyphis spp.), and the

bull shark, Carcharhinus leucas, penetrates more than 4000

km up the Amazon River, reaching Peru. New species have

been described in recent years, particularly of catsharks (e.g.,

Nakaya and Sйret 1999, Last 1999), and additional new species

await formal description (Last and Stevens 1994). Phylogenetic

relationships among ground sharks requires

further study (Naylor 1992), which may eventually result

in the merging of several currently monotypic genera and

some of the families. Compagno (1988) presents a comprehensive

review of the classification and morphology of

Carcharhiniformes.

Squalomorphs (equivalent to the Squalea of Shirai 1992)

are a very diverse and morphologically heterogeneous group

that includes the six- and seven-gill sharks (Hexanchiformes),

bramble sharks (Echinorhiniformes), dogfishes and allies

(Squaliformes), angelsharks (Squatiniformes), sawsharks

(Pristiophoriformes), and rays (Batoidea; fig. 24.2). These

taxa share complete precaudal hemal arches in the tail region,

among many other features (Shirai, 1992, 1996,

Carvalho 1996). Many previous authors defended similar

arrangements for the squalomorphs, but usually excluded

one group or another (e.g., Woodward 1889, White 1937,

Glickman 1967, Maisey 1980). The most dramatic evolutionary

transition among elasmobranchs has taken place within

the squalomorphs—the evolution of rays from sharklike

ancestors, which probably took place in the Early Jurassic

(some 200 Mya). Protospinax, from the Late Jurassic (150

Mya) Solnhofen limestones of Germany, is an early descendent

of the shark–ray transition because it is the most basal

hypnosqualean (fig. 24.2), sister group to the node uniting

angelsharks, sawsharks, and batoids, and has features intermediate

between sharks and rays (Carvalho and Maisey 1996).

Basal squalomorph lineages are relatively depauperate;

hexanchiforms (four genera, five spp.) and bramble sharks

(Echinorhinus, two spp.) are mostly deep-water inhabitants

of the continental slopes but occasionally venture into shallow

water. All species are aplacentally viviparous. Hexanchiforms

have a remarkable longevity; fossil skeletons date from

414 The Relationships of Animals: Deuterostomes

the Late Jurassic. They are united by several derived characters,

such as an extra gill arch and pectoral propterygium

separated from its corresponding radials (Compagno 1977,

Carvalho 1996; compare Shirai 1992, 1996, which do not

support hexanchiform monophyly). The frilled shark,

Chlamydoselachus anguineus, is one of the strangest living

sharks, with an enormous gape, triple-cusped teeth, and eellike

body. Some researchers even thought it was a relic of

Paleozoic “cladodont” sharks (reviewed in Gudger and Smith

1933). Echinorhinus has traditionally been classified with the

Squaliformes (Bigelow and Schroeder 1948, Compagno

1984a) but was given ordinal status by Shirai (1992, 1996,

Carvalho 1996); studies of its dentition further support this

conclusion (Pfeil 1983, Herman et al. 1989).

Squaliformes (20 genera, 121+ spp.), Squatiniformes

(Squatina, 15+ spp.), and Pristiophoriformes (two genera, five

or more spp.) form successive sister groups to the rays

(Batoidea, 73+ genera, 555+ spp.). The squaliform dogfishes

are mesopelagic, demersal, and deep-water species that vary

greatly in size (from 25 cm Euprotomicrus to 6 m Somniosus).

Many species are economically important, and new species

continue to be described (Last et al. 2002). They are aplacentally

viviparous, and some have the longest gestation

periods of all vertebrates (Squalus, some 24 months). Shirai

(1992, 1996) and Carvalho (1996) disagree in relation to the

composition of this order, which is recognized as monophyletic

by Carvalho, but broken into several lineages by Shirai.

Squatiniforms (angelsharks) are morphologically unique,

benthic sharks that resemble rays in being dorsoventrally

flattened with expanded pectoral fins. They are distributed

worldwide, but most species are geographically restricted

(Compagno 1984a). Pristiophoriforms (sawsharks) are poorly

known benthic inhabitants of the outer continental shelves

(Compagno 1984a). They first appear in the fossil record during

the Late Cretaceous of Lebanon (some 90 Mya) and have

an elongated rostral blade (“saw”) with acute lateral rostral

spines that are replaced continuously through life; the saw is

used to stun and kill fishes by slashing it from side to side.

Similar to angelsharks, sawsharks are yolk-sac viviparous.

Rays (batoids), once thought to represent a gargantuan

evolutionary leap from sharklike ancestors (e.g., Regan 1906),

are best understood as having evolved through stepwise anatomical

transformations from within squalomorphs. Sawsharks

are their sister group, sharing with rays various

characters (Shirai 1992), such as enlarged supraneurals

extending forward to the abdominal area. But at least one

feature traditionally considered unique to rays (the antorbital

cartilage) can be traced down the tree to basal squalomorphs,

in the form of the ectethmoid process (Carvalho

and Maisey 1996) of hexanchiforms, Echinorhinus, and squaliforms,

or as an unchondrified “antorbital” in pristiophoriforms

(Holmgren 1941, Carvalho 1996). Even though

“advanced” rays are very modified (e.g., Manta), basal rays

retain various sharklike traits such as elongated, muscular

tails with dorsal fins.

In precladistic days, Batoidea were traditionally divided

into five orders (e.g., Compagno 1977): Pristiformes (sawfishes,

two genera, five or more spp.), “Rhinobatiformes”

(guitarfishes, nine genera, 50+ spp.), Rajiformes (skates, 28

genera, 260+ spp.), Torpediniformes (electric rays, 10 genera,

55+ spp.), and Myliobatiformes (stingrays, 24 genera,

185+ spp.). Phylogenetic analyses have revealed that Rhinobatiformes

is not monophyletic (Nishida 1990, McEachran

et al. 1996), but all other groups are morphologically well

defined (Compagno 1977, McEachran et al. 1996). There is

conflict as to which batoid order is the most basal, whether

it is sawfishes (Compagno 1973, Heemstra and Smith 1980,

Nishida 1990, Shirai 1996) or electric rays (Compagno 1977,

McEachran et al. 1996). The most comprehensive phylogenetic

study to date is that of McEachran et al. (1996);

molecular analyses have hitherto contributed very little

to the resolution of this problem (e.g., Chang et al. 1995).

Rays are clearly monophyletic, with ventral gill openings,

synarcual cartilages, and an anteriorly expanded propterygium,

among other characters (e.g., Compagno 1973,

1977). There is as much morphological distinctiveness

among the different groups of rays as there is among the

orders of sharks. The oldest ray skeletons are from the Late

Jurassic of Europe and are morphologically reminiscent of

modern guitarfishes (Saint-Seine 1949, Cavin et al. 1995),

but their relationships require further study (see Carvalho,

in press).

Sawfishes are large batoids (up to 6 m long), present in

inshore seas and bays, but also in freshwaters. The precise

number of species is difficult to determine because of the

paucity of specimens but is between four and seven; some

are critically endangered because of overfishing and habitat

degradation (Compagno and Cook 1995). They differ from

sawsharks in the arrangement of canals for vessels and nerves

within the rostral saw and in the mode of attachment of rostral

spines. Guitarfishes are widespread in tropical and warm

temperate waters, and are economically important. Much

work is needed on their species level taxonomy; the last

comprehensive revision was by Norman (1926). Characters

supporting their monophyly are known, but they are

undoubtedly a heterogeneous assemblage that requires subdivision

(as in McEachran et al. 1996); for simplicity they are

treated as a single taxon in figure 24.2. Electric rays are notorious

for their electrogenic abilities. Although known since

antiquity, they have been neglected taxonomically until very

recently (e.g., Carvalho 1999, 2001). Their electric organs

are derived from pectoral muscles and can produce strong

shocks that are actively used to hunt prey (Bigelow and

Schroeder 1953, Lowe et al. 1994). All electric ray species

are marine, in tropical to temperate waters, and some occur

in deep water. Skates are oviparous (all other rays are viviparous),

marine, mostly deep water and more abundant in temperate

areas. They also produce weak discharges from caudal

electric organs (Jacob et al. 1994). Even though skates are

the most species-rich chondrichthyan assemblage, they are

Gnathostome Fishes 415

rather conservative morphologically. Rajiform intrarelationships

have been studied by McEachran (1984), McEachran

and Miyake (1990), and McEachran and Dunn (1998). Many

new species still await description (J. D. McEachran, pers.

comm.). Stingrays are also highly diverse (Last and Stevens

1994) and are found in both marine and freshwaters (the 20+

species of South American potamotrygonid stingrays are the

only supraspecific chondrichthyan group restricted to freshwater).

Stingrays can be very colorful and range from 15 cm

(Urotrygon microphthalmum) to 5 m (Manta) across the disk.

Stingray intrarelationships have been recently investigated

by Nishida (1990), Lovejoy (1996), and McEachran et al.

(1996). Stingray embryos are nourished in utero by milk-like

secretions from trophonemata (Hamlett and Koob 1999);

there are at least 10 undescribed species.

Holocephalans (Chimaeras)

Living holocephalans represent only a fraction of their previous

(mostly Carboniferous) diversity. As a result, fossil

holocephalans (summarized in Stahl 1999) have received

more attention from systematists than have extant forms. The

single surviving holocephalan order (Chimaeriformes) contains

three extant families: Chimaeridae (2 genera, 24+ spp.),

Callorhynchidae (Callorhinchus, three spp.), and Rhinochimaeridae

(three genera, eight spp.). Chimaeras are easily distinguished

from elasmobranchs, with opercular gill covers,

open lateral-line canals, three pairs of crushing tooth plates

with hypermineralized pads (tritors), and frontal tenacula

on their foreheads (Didier 1995). Most species are poorly

known, deep-water forms of relatively little economic significance.

All chimaeras are oviparous, and some of their egg

capsules are highly sculptured (Dean 1906). Relationships

among living holocephalans is summarized by Didier (1995).

New species are still being described (e.g., Didier and Sйret

2002), but relationships among chimaeriform species are

unknown.

Osteichthyes (Bony Fishes)

Before the advent of Phylogenetic Systematics (Hennig 1950,

1966, and numerous subsequent authors), Osteichthyes

constituted only bony fishes; tetrapod vertebrates were classified

apart as coordinate groups (usually ranked as classes).

With the recognition that vertebrate classifications should

strictly reflect evolutionary relationships, it has become

apparent that Osteichthyes cannot include only the bony

fishes, but must also include the tetrapods. Thus, there are

two great osteichthyan groups of approximately equal size:

Sarcopterygii (lobefins and tetrapods) and Actinopterygii

(rayfins). Here, we briefly review the so-called “piscine sarcopterygians,”

or lobefins, before considering the largest,

and most diverse radiation of the jawed fishes, the actinopterygians

or rayfins.

Sarcopterygii (Lobefin Fishes and Tetrapods)

The lobefin fishes and tetrapods comprise some 24,000+ living

species of fishes, amphibians, and amniote vertebrates

(mammals; birds, crocodiles; turtles; snakes, lizards, and kin)

with a fossil record extending to the Upper Silurian. All sarcopterygians

are characterized by the evolutionary innovation

of having the pectoral fins articulating with the shoulder

girdle by a single element, known as the humerus in tetrapods.

In contrast, actinopterygian fishes retain a primitive

condition similar to that seen in sharks, in which numerous

elements connect the fin with the girdle. A rich record of fossil

lobefin fishes provides numerous “transitional forms” leading

to Tetrapoda (Cloutier and Ahlberg 1996, Zhu and

Schultze 1997, Zhu et al. 1999, Clack 2002). Two living

groups survive, lungfishes and coelacanths.

Lungfishes

There are six living species of lungfishes, one in Australia

(Neoceratodus forsteri), one in South America (Lepidosiren

paradoxa), and four in Africa (Protopterus spp.). All are freshwater,

but there are more than 60 described fossil genera

dating back to the Devonian, almost all of which were marine.

Of the living lungfishes all except the Australian species

share an ability to survive desiccation by aestivating in

burrows. This lifestyle is ancient; Permian lungfishes are

commonly found preserved in their burrows. Considerable

controversy surrounds the interrelationship of lungfishes.

Most recent studies place them at (Zhu and Schultze 1997)

or near (Cloutier and Ahlberg 1996) the base of the sarcopterygian

tree, although some ichthyologists have claimed

that they are the closest relatives of Tetrapoda (Rosen et al.

1981), a view recently supported with molecular evidence

by Venkatesh et al. (2001).

Coelacanths

Coelacanths were once thought to have become extinct in

the Cretaceous. The discovery of a living coelacanth off the

coast of South Africa in 1938 caused a sensation in the zoological

community [Weinberg (2000) presents a very readable

history; see also Forey (1998)]. Between the 1950s and

the 1990s, extant coelacanths were thought to be endemic

to the Comoro Islands. But in 1997 Arnaz and Mark Erdmann

photographed a specimen in a fish market in Indonesia

(Sulawesi) and eventually obtained a specimen through

local fishermen (Erdmann 1999). Since that time, coelacanths

have been discovered off South Africa, Kenya, and Madagascar

[see Third Wave Media Inc. (2003) for accounts of these

discoveries and other coelacanth news]. Like lungfishes, the

phylogenetic position of coelacanths has been subject to some

dispute. Cloutier and Ahlberg (1996) placed them at the base

of Sarcopterygii; Zhu and Schultze (1997) placed them near

the clade containing Tetrapoda.

416 The Relationships of Animals: Deuterostomes

Actinopterygii (Rayfin Fishes)

The actinopterygian fossil record is rich, but unlike that of

most other vertebrate groups, there are far more living forms

than known fossils. The exact number of rayfin fishes remains

to be determined, but most authors agree that the group

minimally consists of some 23,600–26,500 living species,

with approximately 200–250 new species being described

each year (Eschmeyer 1998). Early actinopterygian fishes

are characterized by several evolutionary innovations

(synapomorphies) still found in extant relatives (Schultze

and Cumbaa 2001). These include several technical features

of the skull and paired fins, and the composition and morphology

of the scales [see Janvier (1996) for an excellent overview

of actinopterygian anatomy]. The earliest well-preserved

actinopterygian, Dialipina, from the lower Devonian of

Canada and Siberia, retains several primitive features of their

osteichthyian and gnathostome ancestors, such as two dorsal

fins (Schultze and Cumbaa 2001).

Living actinopterygian diversity resides mostly in the

crown group Teleostei (see below), but between the speciesrich

teleosts and the base of Actinopterygii are a number of

small but interesting living groups allied with a much more

diverse but extinct fauna. For example, an actinopterygian

thought to represent the closest living relative of teleost fishes

is the North American bowfin, Amia calva (Patterson 1973,

Wiley 1976, Grande and Bemis 1998). The bowfin is the last

remaining survivor of a much larger group of fishes (the

Halecomorphi) that radiated extensively in the Mesozoic and

whose fossil representatives have been found in marine and

freshwater sediments worldwide. As another example, between

and below the branches leading to the living bichirs

and the living sturgeons and paddlefishes are a whole series

of Paleozoic fishes generally termed “palaeoniscoids.” They

display a dazzling array of morphologies, many paralleling

the body forms now observed among teleost fishes and probably

reflecting similar life styles. A review of this fossil diversity

is beyond the scope of this chapter, but the reader can

refer to Grande (1998) and Gardiner and Schaeffer (1989).

However, fossil diversity has important consequences for our

study of the evolution of characters. When we only consider

living groups on the Tree of Life, we might get the impression

that the appearance of some groups was accompanied

by massive morphological change. This is usually not the

case. When the fossils are included, we gain a very different

impression: most of the evolutionary innovations we associate

with major groups are gained over many speciation

events, and the distinctive nature of the living members of

the group is largely due to the extinction of its more basal

members. Thus, it is true that the living teleost fishes are

distinguished from their closest relatives by a large number

of evolutionary innovations (DePinna 1996). Yet, when we

include all the fossil diversity, this impressive number is,

according to Arratia (1999), significantly reduced. Of course,

this is to be expected; evolution by large saltatory steps is

more the exception than the rule, because derived characters

were acquired gradually. Another example is that

gnathostomes, today remarkably diverse and divergent in

anatomy, appear to have been very similar to each other

shortly after their initial separation, because many features

were primitively retained in now extinct stem gnathostome

lineages (Basden et al. 2000, Maisey and Anderson 2001, Zhu

et al. 2001, Zhu and Schultze 2001).

Living Actinopterygian Diversity

and Basal Relationships

Wiley (1998) and Stiassny (2002) provide nontechnical overviews

of basal actinopterygian diversity, and the review of

Lauder and Liem (1983) remains a valuable and highly readable

summary of actinopterygian relationships. The most

basal of living actinopterygians are the bichirs (Polypteridae),

a small group (11 spp.) of African fishes previously thought

to be related to the lobefin fishes (sarcopterygians), or to form

a third group. Despite past controversy, two recent molecular

studies provide additional support for the birchirs as the

basal living actinopterygian lineage (Venkatesh et al. 2001,

Inoue et al. 2002), and this placement now seems well established.

Compared with other rayfin fishes, birchirs are

distinctive in having a rather broad fin base (even giving the

external appearance of a lobe fin), a dorsal fin composed of

a series of finlets running atop an elongate body, and only

four gill arches. Although the analysis by Schultze and

Cumbaa (2001) places them one branch above the basal

Dialipina, their fossil record only just extends to the Lower

Cretaceous (Dutheil 1999), a geologic enigma, but such a

disparity between the phylogenetic age of a taxon and its first

known fossil occurrence is not uncommon among rayfin

fishes (fig. 24.1).

The living chondrostean fishes include the sturgeons of

the Holarctic and the North American and Chinese paddlefishes.

The comprehensive morphological analyses of Grande

and Bemis (1991, 1996) have established a hypothesis of

relationships among the living and fossils members of this

group, which originated in the Paleozoic. The diversification

of the living chondrosteans may go back to the Jurassic (Zhu

1992), when paddlefishes and sturgeons were already diversified.

Paddlefishes and sturgeons retain many primitive characters,

such as a strongly heterocercal tail that led some 19th

century ichthyologists to believe that they are related to

sharks. Sturgeons are among the most endangered, sought

after, and largest of freshwater fishes. The Asian beluga Huso

huso reaches at least 4 m in length, and a large female may

yield 180 kg of highly prized caviar. Paddlefish caviar is also

prized, and the highly endangered Chinese paddlefish grows

to twice the size of its American cousins, reaching 3 m.

The remaining rayfin fishes belong to the clade Neopterygii.

Garfishes (Lepisosteidae) are considered by most to be the

Gnathostome Fishes 417

basal group (Patterson 1973, Wiley 1976). They form an

exception among rayfin fishes in that there are as many living

gars (a mere seven species) as fossil forms. Although fossils

are known from many regions of the world and their

record extends to the Lower Cretaceous, living gars are now

confined to North and Middle America and Cuba.

Amia calva, the North American bowfin, is the sole living

representative of Halecomorphi, a group that radiated in

the Mesozoic. It shares a number of evolutionary innovations

with teleost fishes (first detailed by Patterson 1973) but also

displays a number of teleost characters that are now considered

convergent, such as having cycloid rather than ganoid

scales. Although most workers have followed Patterson (1973)

in the recognition of Amia as the closest living relative of the

Teleostei, there remains some controversy about their systematic

position (Patterson 1994); alternative schemes of

basal neopterygian relationships and the proximate relatives

of the Teleostei are reviewed in Arratia (2001).

Teleostei

Among vertebrates, without doubt, Teleostei dominate the

waters of the planet. The earliest representatives of living

teleost lineages (the Teleocephala of DePinna 1996) date to

the Late Jurassic some 150 Mya, but as noted by Arratia

(2001), if definitions of the group are to include related fossil

lineages, this date is pushed back into the Late Triassic–Early

Jurassic (~200–210 Mya). Regardless of how fossil lineages

are incorporated into definitions of the group, today’s teleosts

occupy almost every conceivable aquatic habitat from

high-elevation mountain springs more than 5000 m above

sea level to the ocean abyss almost 8500 m below. Estimates

of the number of living species vary, but most authors agree

that a figure of around 26,000 is reasonable. Although discovery

rates are more or less constant at around 200–250

new species a year, for some groups, particularly those in little

explored or inaccessible habitats, new species are being described

in extraordinary numbers, for example, 30 new snailfishes

from deep water off Australia (Stein et al. 2001) with

some 70 more to be described from polar seas, or an estimated

200 new rock-dwelling cichlids from Lake Victoria,

Africa (Seehausen 1996). There are more teleost species than

all other vertebrates combined, and their number contrasts

starkly with the low species diversity in their immediate

amiiform relatives, or indeed of all basal actinopterygian lineages.

Among actinopterygians the extraordinary species

richness of the teleostean lineage is noteworthy, and although

“adaptationist” explanations are not readily testable, it seems

probable that much of their success may be attributed to the

evolution of the teleost caudal skeleton, permitting increased

efficiency and flexibility in movement (Lauder 2000), and to

the evolution of powerful suction feeding capabilities that

have facilitated a wide range of feeding adaptations (Liem

1990).

Teleostean Basal Relationships

Systematic ichthyology has a rich history, and the past three

centuries have seen waves of progress and revision. But in the

modern era, perhaps one of the most important contributions

on teleost relationships was that of Greenwood et al. (1966;

fig. 24.3). In that paper, the authors presented a tentative

scheme of relationships among three main lineages,

Elopomorpha (tarpons and eels), Osteoglossomorpha (elephantfishes

and kin), and what are now known as the

Euteleostei (all “higher teleosts,” including such groups as cods

and basses). Greenwood et al. (1966) found placement of

Clupeomorpha (herrings and allies) problematic, but most

subsequent workers have placed them as the basal euteleosts.

Recently, however, this alignment has been challenged (see

below). As Patterson (1994) later noted, it was as if the distinction

between monotremes, marsupials, and placental

mammals was not recognized until the mid 1960s.

By 1989, Gareth Nelson summarized the previous 20

years of ichthyological endeavor with the by now much

quoted observation that “recent work has resolved the bush

at the bottom but that the bush at the top persists.” He presented

a summary tree that showed a fully resolved scheme

of major teleostean lineages as a comb leading to the spiny

rayed Acanthomorpha that contains the percomorph “bush

at the top.”

The outstanding problem of Percomorpha is discussed

below, but it is perhaps also worth noting that some recent

studies have begun to challenge the notion of a fully resolved

teleostean tree and to question the monophyly of some lineages

(e.g., Lк et al. 1993, Johnson and Patterson 1996,

Arratia 1997, 1999, 2000, 2001, Filleul and Lavouй 2001,

Inoue et al. 2001, Miya et al. 2001, 2003). This is perhaps

not surprising given that Nelson (1989) was somewhat

guarded in his optimism and noted that although the interrelationships

of major groups of fishes were resolved no

group was defined by more than a few characters. Results of

more refined matrix-based analyses that incorporate broader

taxon sampling than the previously more standard “exemplar

“ approaches, the inclusion of new high quality fossil

data, and the beginnings of more sophisticated multigene

molecular studies indicate that character support for many

teleost nodes is weak, ambiguous, or entirely wanting. Some

of these changes or uncertainties are reflected in figure 24.1,

in which basal teleostean relationships are represented

as unresolved. For example, in a highly influential paper,

Patterson and Rosen (1977) hypothesized that osteoglossomorphs

are the sister group of elopomorphs and other

living teleosts, whereas Shen (1996) and Arratia (e.g., 1997,

1999) have proposed that elopomorphs occupy that basal

position.

We turn now to a brief review of diversity within extant

non-acanthomorph teleost groups. Osteoglossomorpha consist

of two freshwater orders: the North American Hiodonti418

The Relationships of Animals: Deuterostomes

formes (mooneyes; two spp., one family) and mostly Old

World Osteoglossiformes (bony tongues, knifefishes, and

elephantfishes; 220+ spp., five families). Osteoglossomorpha

are an ancient group with a long fossil history dating to the

Jurassic (Patterson 1993, 1994, Li and Wilson 1996) and

displaying a number of primitive characters as well as two

evolutionary innovations; a complex tongue-bite mechanism

and a gut that uniquely coils to the left of the stomach. The

most speciose and perhaps the most interesting members of

this group are the elephantfishes (Mormyridae), which create

an electric field with muscles of the caudal region and

use it to find prey and avoid obstacles in their turbid water

habitats. Relationships among mormyrids and the evolution

of their electric organs have recently been elucidated with

molecular data by Sullivan et al. (2000) and Lavouй et al.

(2000). Other osteoglossiforms include the large (to 2.5 m)

bonytongues of South America, Asia, and Africa. Li and

Wilson (1996) analyzed phylogenetic relationships and discussed

evolutionary innovations of osteoglossomorphs, and

a recent molecular analysis (Kumazawa and Nishida 2000)

corroborates osteoglossomorph monophyly but differs in its

assessment of osteoglossiform interrelationships.

Elopomorpha are a heterogeneous group united by the

unique, leaflike, transparent leptocephalus larval stage, once

considered a distinct taxonomic group, and by the possession

of derived sperm morphology (Mattei 1991, Jamieson

1991). All are marine, although some venture into brackish

waters. Elopomorph intrarelationships are poorly understood;

however, most studies agree in placing Elopiformes

(tarpons and ladyfishes; eight spp., two families) as the basal

order. Albuliformes (bonefishes, two spp., one family) are a

small group highly prized by fishermen. Notacanthiformes

(halosaurs and spiny eels, 25 spp., two families) are marine,

deep-water fishes. The bulk of elopomorph diversity lies in

the Anguilliformes (true eels, 750+ spp., 15 families), which

includes morays (200 spp.), snake eels (250 spp.), conger

eels (150 spp.), and the anadromous freshwater eels (15

spp.). Saccopharyngiformes (deep-water gulper eels, 25 spp.,

three families) contains among the most bizarre of living

vertebrates, with luminescent organs and huge mouths capable

of swallowing prey several times their body size. Forey

et al. (1996) accepted elopomorph monophyly and presented

a detailed study of their intrarelationships, using both morphological

and molecular characters. However, two recent

studies (Filleul and Lavouй 2001, Obermiller and Pfeiler

2003) have challenged elopomorph monophyly, and Filleul

and Lavouй (2001) place the four orders as incertae sedis

among basal teleosts.

Figure 24.3. Diagram of teleostean relationships from

Greenwood et al. (1966). This remarkably prescient,

precladistic study delineated for the first time the major

groups of teleostean fishes and thereby laid an important

foundation for the “modern era” of teleostean systematics

that was to follow.

Gnathostome Fishes 419

Until 1996, the remaining teleost fishes were grouped

into two putative lineages, Clupeomorpha (herrings and allies,

360+ spp., five families) and Euteleostei. Euteleostei have

proven to be a problematic group, persistently defying unambiguous

diagnosis (Fink 1984).

Following the molecular work of Lк et al. (1993), Lecointre

(1995) and Lecointre and Nelson (1996) suggested, based on

both morphological and molecular characters, that ostariophysans

(minnows, catfishes, and allies) are not euteleosts

but instead are the sister group of clupeomorphs. Further

evidence is emerging, both molecular (Filleul and Lavouй

2001, G. Orti pers. comm.) and morphological (Arratia 1997,

1999, M. DePinna pers. comm.) to support this hypothesis,

which removes one of the stumbling blocks to understanding

the evolution of euteleosts, but its validity and implications

are not yet fully understood. For example, Ishiguro et al. (2003)

find mitogenomic support for an Ostariophysan-clupeomorph

clade, but one that also includes the alepocephaloids (slickheads,

see below) nested within it.

With the ostariophysans removed, Johnson and Patterson

(1996) argued that four unique evolutionary innovations characterize

the “new” Euteleostei and recognized two major lineages.

The first, Protacanthopterygii, is a refinement of the

group first proposed by Greenwood et al. (1966). The second

(Neognathi) placed the small order Esociformes (the freshwater

Holarctic pikes and mudminnows; about 10 spp., two families)

as the sister group of the remaining teleosts (Neoteleostei).

The relationships of the pikes and mudminnows remain problematic,

but they share two unique evolutionary innovations

with neoteleosts (Johnson and Patterson 1996).

The reconstituted Protacanthopterygii consists of two

orders, Salmoniformes and Argentiniformes, each with two

suborders. Salmoniformes includes the whitefishes, Holarctic

salmons and trouts, Salmonoidei (65+ spp., one family)

and the northern smelts, noodlefishes, southern smelts and

allies, and Osmeroidei (75+ spp., three families). The

Argentiniformes include the marine herring smelts and allies

(Argentinoidei; 60+ spp., four families), most of which

occur in deep water, and the deep-sea slickheads and allies

(Alepocephaloidea; 100+ spp., three families).

Morphological character support for a monophyletic Neoteleostei

and the monophyly and sequential relationships of

the three major neoteleost groups leading to Acanthomorpha,

depicted in figure 24.1, appears strong (Johnson 1992,

Johnson and Patterson 1993, Stiassny 1986, 1996), and it is

perhaps at this level on the teleostean tree that most confidence

can currently be placed. Stomiiformes (320+ spp., four

families) are a group of luminescent, deep-sea fishes with

exotic names such as bristlemouths and dragonfishes that

complement their morphological diversity (fig. 24.4). Two

genera of midwater bristlemouths (Cyclothone and Gonostoma)

have the greatest abundance of individuals of any

vertebrate genus on Earth (Marshall 1979). Harold and Weitzman

(1996) provide the most recent analysis of stomiiform

intrarelationships. Aulopiformes (220+ spp., 15 families) are

a diverse group of nearshore and mostly deep-sea species,

including the abyssal plain tripod fishes, the familiar tropical

and temperate lizardfishes, and midwater predators such

as the sabertooths and lancetfishes (for the most recent analyses

of their intrarelationships, see Johnson et al. 1996,

Baldwin and Johnson 1996, Sato and Nakabo 2002). Members

of Myctophiformes—lanternfishes and allies (240+ spp.,

two families)—are also ubiquitous midwater fishes, most

with luminescent organs. They are a major food source for

economically important midwater feeders, from tunas to

whales, and many undertake vertical migrations into surface

waters at night to feed, returning to depths during the day,

thereby contributing significantly to biological nutrient cycling

in the deep ocean. Stiassny (1996) and Yamaguchi

(2000) provide recent analyses of their intrarelationships.

Acanthomorpha and the “Bush at the Top”

The spiny-rayed fishes, Acanthomorpha, are the crown group

of Teleostei. With more than 300 families and approximately

16,000 species, they comprise more than 60% of extant teleosts

and about one-third of all living vertebrates. This immense

group of fishes exhibits staggering diversity in adult

and larval body form, skeletal and soft anatomy, size (8 mm

to 15 m), habitat, physiology, and behavior. Acanthomorphs

first appear in the fossil record at the base of the Late Cretaceous

(Cenomanian) represented by more than 20 genera

assignable to four or five extant taxa (fig. 24.1). By the late

Paleocene the fauna is somewhat more diverse, but at the

Middle Eocene, as seen in the Monte Bolca Fauna, an explosive

radiation seems to have occurred, wherein the majority

of higher acanthomorph diversity is laid out (Patterson 1994,

Bellwood 1996). To date, because of the uncertainty of structure

and relationships of many of the earlier fossils and the

rapid appearance of most extant families, fossils have offered

little to our understanding of acanthomorph relationships.

Acanthomorpha originated with Rosen’s (1973) seminal

paper on interrelationships of higher euteleosts and was

based on five ambiguously distributed characters. In an attempt

to define the largest and most diverse acanthomorph

assemblage, Percomorpha, Johnson and Patterson (1993)

proposed a morphology-based hypothesis of acanthomorph

relationships. In so doing, they reviewed and evaluated support

for previous hypotheses, including acanthomorph

monophyly, for which they identified eight evolutionary innovations.

Perhaps the most convincing of these are the presence

in the dorsal and anal fins of true fin spines, as well as

a single median chondrified rostral cartilage associated with

specific rostral ligaments (Hartel and Stiassny 1986, Stiassny

1986) that permit the jaws to be greatly protruded while

feeding. Johnson and Patterson (1993) proposed a phylogeny

for six basal acanthomorph groups leading sequentially

to a newly defined Percomorpha. Below, we briefly discuss

acanthomorph diversity in this proposed phylogenetic order

(fig. 24.5).

420 The Relationships of Animals: Deuterostomes

Interestingly, Lampridiformes (opahs and allies) were

once placed among the perciform fishes at the top of the tree.

They are a small (20 spp., seven families) but diverse group,

characterized by a uniquely configured, highly protrusible

upper jaw mechanism. Except for the most primitive family,

the velifers, which occur in near shore-waters, the remaining

families are meso- and epipelagic. In body shape they

range from the deep-bodied opahs to extremely elongate

forms such as the oarfish (Regalecus glesne), which is the longest

known bony fish, reported to reach 15 m. The position

of lampridiforms as a basal acanthomorph group has been

supported by both morphological (Olney et al. 1993) and

molecular data sets (Wiley et al. 2000, Miya et al. 2001, 2003,

Chen et al. 2003).

Polymixiiformes (beard fishes; 10 spp., one family) are

characterized by two chin barbels supported by the first

branchiostegals and occur on the continental shelf and upper

slope. The fossil record for this group is considerably

more diverse than its living representation. Recent molecular

studies have confirmed a basal position for these fishes,

but some suggest a placement within a large clade consisting

otherwise of paracanthopterygian and zeoid lineages (e.g.,

Miya et al. 2001, 2003, Chen et al. 2003).

Paracanthopterygii (1,200+ spp., 37 families) are an odd

and almost certainly unnatural assemblage of freshwater and

marine fishes first proposed by Greenwood et al. (1966) and

refined to its present form by Patterson and Rosen (1989).

Most of the hypothesized evolutionary innovations proposed

by these authors are suspect (Gill 1996), and molecular studies

by Wiley et al. (2000) and Miya et al. (2001) suggest that

although the freshwater Percopsiformes (troutperches; six

spp., three families) and Gadiformes (cods; 500+ spp., nine

families) are basal acanthomorphs, the other groups may be

scattered through the higher acanthomorph lineages. These

orders include Ophidiiformes (cuskeels; 380+ spp., 18 families),

Batrachoidiformes (toadfishes; 70 spp., three families),

and Lophiiformes (anglerfishes; 300+ spp., 18 families). Most

species belonging to these orders are marine. The dismemberment

of all or part of Paracanthopterygii will have significant

implications for acanthomorph relationships, perhaps

particularly those within the perciforms.

Between the paracanthopterygians and the immense diversity

of Percomorpha are three small, but phylogenetically

critical, marine lineages. Stephanoberyciformes (90 spp., nine

families) is a monophyletic group of marine benthic and

deep-water fishes commonly called pricklefishes and whalefishes.

Johnson and Patterson (1996) separated this group

from Beryciformes, but molecular data suggest that at least

some members of the group might rejoin Beryciformes (Wiley

et al. 2000, Colgan et al. 2000, Chen et al. 2003). Zeiformes

(45 spp., five families) includes the dories, a marine group

of deep-bodied fishes that includes the much-valued John

Figure 24.4. The viperfish,

Chauliodus sloani; anatomical

detail from Tchernavin (1953).

Larvae redrawn after Kawaguchi

and Moser (1984). Teleostean

fishes are biomechanically

complex; the head alone is

controlled by some 50 muscles

operating more than 30

movable skeletal parts. Such

anatomical complexity, plus a

wide range of ontogenetic

variation, ensures a continued

pivotal role for anatomical input

into systematic study.

Gnathostome Fishes 421

Dory of the Atlantic. Recent molecular studies suggest a relationship

between the dories and the codfishes and/or beardfishes

(Wiley et al. 2000, Miya et al. 2001, Chen et al. 2003),

but this conclusion might be due to the relatively low numbers

of species included in these studies. Beryciformes (140+

spp., seven families) includes some of the most familiar reefdwelling

fishes, the squirrelfishes. Beryciforms are entirely

marine and occur worldwide from shallow depths, where

they are nocturnal, to the deep sea. External bacterial luminescent

organs characterize the pinecone fishes and flashlight

fishes, the latter having a complex mechanism for rapidly

occluding the large subocular light organ by rotating it downward

or covering it with a lidlike shutter. Two genera of the

closely related roughies (Trachichthyidae) have internal

luminescent organs, and the orange roughy (Hoplostethus

atlanticus) is an overexploited food fish.

Percomorpha, the Bush at the Top

Percomorph (14,000+ spp., 244 families) are the crown group

of the spiny-rayed fishes and best represent what Nelson

(1989) called the “bush at the top.” The name Percomorpha

originated with Rosen (1973) and was essentially the equivalent

of Greenwood et al.’s (1966) Acanthopterygii, which consisted

of beryciforms, perciforms, and groups placed between

and beyond those two, such as lampridiforms, zeiforms, gasterosteiforms,

scorpaeniforms, pleuronectiforms, and tetraodontiforms.

Rosen presented no characters in support of his

Percomorpha, nor have any been supported subsequently (but

see Stiassny 1990, 1993, Stiassny and Moore 1992, Roberts

1993). The major goal of Johnson and Patterson’s (1993) analysis

was to sort out basal lineages of acanthomorphs and revise

the composition of Percomorpha to represent a monophyl-

Figure 24.5. Intrarelationships

among acanthomorph lineages

after Johnson and Patterson

(1993).

422 The Relationships of Animals: Deuterostomes

etic group diagnosed by derived characters. In the process,

they erected a new, putatively monophyletic assemblage,

Smegmamorpha, which, together with “the perciforms and

their immediate relatives,” constituted the newly defined

Percomorpha. They identified eight evolutionary innovations

of the Percomorpha, all of which are homoplasious. Although

monophyly of Johnson and Patterson’s Percomorpha has not

been challenged subsequently with morphological analyses,

it is considered tenuous, particularly in view of our ignorance

of the composition and intrarelationships of Perciformes and

allies (below) and strong doubts about paracanthopterygian

monophyly. To date, no molecular analyses have captured a

monophyletic Percomorpha without the inclusion of certain

“paracanthopterygian” lineages.

Smegmamorpha (1,700+ spp., 37 families) of Johnson

and Patterson (1993) are a diverse group consisting of spiny

and swamp eels (Synbranchiformes; 90 spp., three families),

gray mullets (Mugiliformes; 80 spp., one family), pygmy

sunfishes (Elassomatiformes; six spp., one family), sticklebacks,

pipefishes and allies (Gasterosteiformes; 275 spp., 11

families), and the speciose silversides, flyingfishes, killifishes,

and allies (Atherinomorpha; 1225+ spp., 21 families, four

orders). The recognition of this group was greeted with some

skepticism because swamp and spiny eels had traditionally

been allied with the perciforms whereas pygmy sunfishes had

been considered centrarchids (sunfish and basses), a family

deeply embedded in one suborder of Perciformes. Smegmamorpha

is united by a single evolutionary innovation,

a specialized attachment of the first intermuscular bone

(epineural) at the tip of a prominent transverse process on

the first vertebra, but several additional specializations are

shared by most smegmamorphs. There have been no comprehensive

morphological analyses to challenge smegmamorph

monophyly; however, Parenti (1993) suggested

that atherinomorphs might be the sister group of paracanthopterygians,

and Parenti and Song (1996) identified a

pattern of innervation of the pelvic fin in mullets and pygmy

sunfishes that is shared with more derived perciforms. Molecular

analyses have failed to capture monophyly of smegmamorphs,

although major components of the group are

recognized (e.g., Wiley et al. 2000, Miya et al. 2003, Chen et al.

2003). Although relationships among smegmamorphs remain

unknown, Stiassny (1993) suggested grey mullets (Mugilidae)

may be most closely related to atherinomorphs, and Johnson

and Springer (1997) presented evidence suggesting a possible

relationship between pygmy sunfishes and sticklebacks.

The remaining groups comprise some 12,000+ species

in more than 207 families. In their cladogram of percomorph

relationships (fig. 24.4), Johnson and Patterson (1993)

placed Perciformes (perches and allies) in an unresolved

polytomy with Smegmamorpha and four remaining groups

traditionally classified as orders: the scorpionfishes and allies

(Scorpaeniformes), flying gurnards (Dactylopteriformes),

flatfishes (Pleuronectiformes), and triggerfishes, pufferfishes,

and allies (Tetraodontiformes). However, they saw no reason

to exclude these last four orders from the traditional

Perciformes and believed it likely that they are nested within

it. Subsequently, Mooi and Gill (1995) classified Scorpaeniformes

within Perciformes. To date, no morphological or

molecular synapomorphies support a monophyletic Perciformes

in either the restricted or expanded sense that would

include any or all of the orders Johnson and Patterson (1993)

placed in their terminal polytomy. Many questions remain

about monophyly and interrelationships of a number of the

approximately 25 suborders and more than 200 families

included in that polytomy. Certainly the possibility that affinities

of some members lie with other acanthomorphs, or

vice versa, cannot be dismissed. With these observations in

mind, we review the remaining orders.

Perciformes (9800+ spp., 163 families) are the largest and

most diverse vertebrate order. Perciforms range in size from

the smallest vertebrate, the 8 mm Trimmatom nanus (for which

an estimated 3674 individuals would be needed to make up

one quarter-pound gobyburger), to the 4.5 m black marlin

(Makaira indica). Although there are a number of freshwater

perciforms (mostly contained within the large cichlid clade),

most species are marine, and they represent the dominant

component of coral reef and inshore fish faunas. In a taxonomic

sense, Perciformes is a catchall assemblage of families and suborders

whose relationships have not been convincingly shown

to lie elsewhere. Although there is reasonably good support

for monophyly of about half of the suborders, others remain

poorly defined, most notably the largest suborder, Percoidei

(3,500+ spp., 70 families), another catch-all or “wastebasket

group,” for which not a single diagnostic character has been

proposed. Percoids are usually referred to as perchlike fishes,

and although this general physiognomy characterizes many

families, such as freshwater perches (Percidae), sunfishes

(Centrarchidae), sea basses (Serranidae), and others, percoids

encompass a wide range of body forms, from the deep-bodied

moonfishes (Menidae), butterflyfishes (Chaetodontidae),

and more, to very elongate, eel-like forms such as bandfishes

(Cepolidae) and bearded snakeblennies (Notograptidae). For

lists and discussions of perciform suborders and percoid families,

see Johnson (1993), Nelson (1994), and Johnson and Gill

(1998), each of which, not surprisingly, differ somewhat in

definition and composition of the two groups.

Scorpaeniformes (lionfishes and allies; 1,200+ spp., 26

families) were included within Perciformes by Mooi and Gill

(1997) based on a specific pattern of epaxial musculature

shared with some perciforms. It is a large, primarily marine

group characterized by the presence of a bony stay of questionable

homology that extends from the third infraorbital

across the cheek to the preopercle. Monophyly, group composition,

and relationships remain controversial, but most

recent work supports two main lineages, scorpaenoids and

cottoids (e.g., Imamura and Shinohara 1998), and preliminary

molecular studies suggest a close relationship between

zoarcoids and the cottoid lineage (Miya et al. 2003, Smith

2002, Chen et al. 2003). Whether the scorpaenoid and cotGnathostome

Fishes 423

toid lineages are sister groups is open to question, and clarification

of scorpaeniform relationships is an important component

of the “percomorph problem.”

Dactylopteriformes (flying gurnards; seven spp., one family)

are a small, clearly monophyletic, group of inshore bottom-

dwelling marine fishes characterized by a thick, bony,

“armored” head with an elongate preopercular spine and

colorful, greatly enlarged, fanlike pectoral fins. Their relationships

are obscure (Imamura 2000), and they have been variously

placed with, among other groups, the scorpaeniforms

and gasterosteiforms. Molecular studies to date have shed

little light on placement, with weak support for an alignment

with flatfishes (Miya et al. 2001), gobioids (Miya et al. 2003),

or syngnathoids (Chen et al. 2003).

Pleuronectiformes (flatfishes; 540+ spp., seven families) are

widely distributed, bottom-dwelling fishes containing a number

of commercially important species. These are characterized

by a unique, complex evolutionary innovation in which

one eye migrates ontogenetically to the opposite side of the

head, so that the transformed juveniles and adults are asymmetrical

and lie, eyeless side down, on the substrate. Their

relationships as shown by morphological analysis have most

recently been reviewed by Chapleau (1993) and Cooper and

Chapleau (1998). A molecular analysis of mitochondrial ribosomal

sequences by Berendzen and Dimmick (2002) suggests

an alternative hypothesis of relationship. Interestingly, a recent

mitogenomic study provides quite strong nodal support

for a relationship with the jacks (Carangidae), but taxon sampling

in this region of the tree is quite sparse (Miya et al. 2003).

Tetraodontiformes (triggerfishes, puffers, and allies; 350+

spp., 10 families) are a highly specialized and diverse order of

primarily marine fishes, ranging in size from the 2 cm diamond

leatherjacket (Rudarius excelsus) to the 3.3 m (>1000 kg) ocean

sunfish (Mola mola). They are characterized by small mouths

with few teeth or teeth incorporated into beaklike jaws, and

scales that are either spine like or, more often, enlarged as plates

or shields covering the body as in the boxfishes (Ostraciidae).

Members of three families have modified stomachs that allow

extreme inflation of the body with water as a defensive mechanism.

Relationships of tetraodontiforms have been treated in

large monographs dealing with comparative myology (Winterbottom

1974) and osteology (Tyler 1980). Although tetraodontiforms

have been considered as highly derived

percomorphs, Rosen (1984) proposed that they are more

closely related to caproids and the apparently more basal

zeiforms. Johnson and Patterson (1993) rejected that hypothesis,

as do ongoing molecular studies (Holcroft 2002, N. I.

Holcroft pers. comm.). However, it is defended in a recent

morphological analysis (Tyler et al. 2003).

Concluding Remarks

Systematic ichthyologists were early to adopt Hennig’s methods

and have made great progress toward understanding the

evolutionary diversification of fishes. Much of the new phylogenetic

structure is underpinned by morphological character

data, most of it from the skeleton and much of it

gathered anew or reexamined and refined during the last 35

years. Another seminal innovation appeared fortuitously on

the cusp of the cladistic revolution—the use of trypsin digestion

in cleared and stained preparations, followed by the

ability to stain cartilage as well as bone. These techniques

revolutionized fish osteology and greatly facilitated detailed

study of skeletal development adding significantly to our

understanding of character transformation and homology.

However, there is still much to do. Our understanding of the

composition and relationships of Percomorpha, with more

than half the diversity of all bonyfishes, remains chaotic—a

state of affairs proportionally equivalent to not knowing the

slightest thing about the relationships among amniote vertebrates.

Fishes are a tremendously diverse group of anatomically

complex organisms (e.g., fig. 24.4) and undoubtedly morphology

will continue to play a central role in systematic

ichthyology. However, as in other groups of organisms, molecular

analyses are increasingly beginning to make significant

contributions, especially for fish groups with confusing

patterns of convergent evolution. The combination of molecular

and morphological data sets, and the reciprocal illumination

they shed, augurs an exciting new phase in

systematic ichthyology. We are, perhaps, at the halfway point

of our journey.

Acknowledgments

We gratefully acknowledge the numerous colleagues whose

studies of fish phylogenetics have helped to elucidate the

present state of the art for the piscine limb of the Tree of Life,

and extend our apologies to those we may have omitted or

inadvertently misrepresented in our efforts to keep this chapter

to a manageable length. Thanks also to Scott Schaefer and Leo

Smith (AMNH) for some helpful comments on an early draft of

the manuscript, and additional thanks to Leo for his artful help

with the figures that accompany the chapter. Part of this work

was funded through grant DEB-9317881 from the National

Science Foundation to E.O.W. and G.D.J. and through the

Scholarly Research Fund of the University of Kansas to E.O.W.

We thank both institutions. Ongoing support from the Axelrod

Research curatorship to M.L.J.S. is also gratefully acknowledged.

Finally our thanks to Joel Cracraft and Mike Donoghue for so

successfully having taken on the formidable task of organizing

the Tree of Life symposium and without whose constant

nudging this chapter would never have seen the light of day.

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