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Family: Amblyopsidae
cavefishes |
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Introduction:
The Percopsiformes
(trout-perches, sand rollers, pirate perchers, and
cavefishes) |
Fig.
1. Comparative external morphology of amblyopsid species (drawing by
John
Ellis; from Romero, 2003b)
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The Amblyopsidae is one of the three families
of the Order Percopsiformes. The oldest fossils of this Order date
back to the Paleocene Paskapoo Formation (between 62 and 60 MYA) in
Western Canada (Murray and Wilson 1996). The Percopsiformes may be
remotely related to the codfishes (Gadiformes) and toadfishes (Batrachoidiformes).
These are fishes small in size (less than 20 cm or 8 in) characterized
by a mosaic of primitive and advanced characters that include some
primitive ones such as adipose fin and advanced ones such as a pelvic
girdle located further back from the cranium when compared with most
other fishes. They also have fewer fin spines and ray-supported
dorsal and anal fins, each usually with one to four anterior soft
spines. If the pelvic fins are present, they are located in a
subabdominal position behind pectorals, with three to eight soft
rays. Their body is covered with cycloid or ctenoid scales.
Percopsiformes are found in North America, from
Alaska and the Great Lakes drainage to the south and eastern United
States. All species are freshwater with two species found in swamps,
one as a facultative cave dweller, and four species are obligatory
cavernicoles. They are opportunistic predators that eat a variety of
food items and at least one species is cannibalistic. Besides all
species being solitary, little is known except for particular
behaviors studied among cave fishes (see below). At least two of the
non-cavernicolous species are nocturnal. They are oviparous but
nothing else is known as to make further generalizations. Spawning (at least for the non-cavernicolous species)
takes place in the Spring. Fecundity tends to be low.
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This order is usually divided into two suborders: 1. Percopsoidei, characterized by the presence of an
adipose fin and a complete lateral line; they are represented by one
family (Percopsidae or trout perches) with one genus and two species;
and, 2. Aphredoderoidei, characterized by the
absence of an adipose
fin and
includes two families: Aphredoderidae (pirate perch, one species) and
Amblyopsidae (swamp and cave fishes, four genera and six species)
(Romero 2003). One of the facts that has been very controversial
about the phylogeny of this order is that they show primitive
conditions such as the presence of an adipose fin, which suggests
character reversal in their evolutionary history. Some have
suggested that the amblyopsids should be placed in a separate
suborder: Amblyopsoidei. The monophyly of
the Percopsiformes has been questioned by Rosen (1985) and Patterson
and Rosen (1989). Murray and Wilson (1999) proposed to remove the
amblyopsid family from it and created a new order: Amblyopsiformes. |
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The Amblyopsidae |
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NON-TROGLOMORPHIC SPECIES
There is only one non-troglomorphic
species of Amblyopsidae |
| The Amblyopsidae is a
freshwater family distributed in the southern and eastern
(unglaciated) United States (Figure 1). They are characterized by
having a large, flattened head, oblique mouth, strongly protruding lower jaw, large branchial cavity (probably for oral incubation), a jugular vent
(anus), and small embedded cycloid scales, except in the head which is
naked. Dorsal and anal fins have similar shapes. They have rows of sensory papillae on the head, body, and
tail. Their eyes range from small (microphthalmic) in the epigean and
troglophilic species to vestigial (remanent eye tissue under the skin)
in the troglomorphic ones. Troglomorphic species are also
characterized for: (a) being depigmented (looking pinkish because of
the blood vessels showing through the translucent skin, with only a
few, mostly non-functional melanophores), (b) low metabolism, and (c)
low fecundity. Because of the jugular position of the genital papilla
and the attachment of the gill membranes to the isthmus, Woods and Inger (1957) proposed that all amblyopsids carry eggs and yolk-sac fry
in their gill cavities; however, such a behavior has been observed
only in the genus Amblyopsis. The systematics of this
family needs revision since genetic studies have shown it to be much
more complex that previously believed (Bergstrom 1997). The six
species amblyopsids represent the transition from epigean to hypogean
waters:
Chologaster cornuta (epigean),
Forbesichthys agassizii (troglophile or facultative
cavernicole) and Typhlichthys
subterraneus, Amblyopsis rosae,
A. spelaea, and
Speoplatyrhinus poulsoni,
all troglomorphic (Figure 2) (Cooper & Kuhene 1974, Page & Burr 1991,
Romero 2001a).
Comparative characteristics are summarized in Table
1 below.
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The swamp
fish, Chologaster cornuta Agassiz, 1853

This is the only species of the amblyopsid family that
is not found in the hypogean environment and, therefore, is considered to
be fully epigean. However, its study is helpful in order to discern
the evolution of the family as a whole. This species shows a stark
contrast between its dorsally brown and ventrally creamy white
colorations. Individuals of this species also show three narrow, dark
stripes on each side, one on lower side wide at front, narrow at rear;
and two black streaks along back before dorsal fin separate to
encircle dorsal fin. They have a dusky black band in the dorsal fin
while the caudal fin is clear near the base, often black in the
center. They have 9 to 12
dorsal
soft rays, 9 to 11 branched caudal rays, and 9 to 10
anal
soft rays. They lack pelvic fins. Other morphological characters
include small eyes, 0 to 2 rows of papillae on caudal fin and visible
pink gills. Males have an appendage on the dorsal surface of the
snout which have been implicated in courtship and in chemoreception,
but neither hypotheses has been verified (Ross and Rohde, 2003). This
is the only amblyopsid species living along the Atlantic Coastal
Plain; it is found from the Roanoke River drainage in southeast
Virginia to Altamaha River drainage in central Georgia (Cooper and
Rohde, 1980; Rohde and Pernell, 1994; Figure 2). Typical habitats are
waters well-shaded by vegetation and debris in swamps, ponds, ditches,
and slow streams (but not in caves) (Shute et al. 1981). Water temperatures where they
are found never exceed 23°C. Although this species is locally common,
it is hard to spot because both being largely nocturnal and found in
heavily vegetated waters (Rohde and Pernell 1994, p. 139; Ross &
Rohde, 2003). They feed mostly at night, on small amphipods, aquatic
insects, cladocerans, ostracods, and copepods. They reach sexual
maturity atve been given to this species: C. cornutus
and C. avitus. one year of age. Spawning takes place in March and April
and they may live up to two years. There is no evidence that they are
gill brooders (Ross and Rohde, 2003). Two synonims have been
given to this species in the past: C. cornutus and C.
avitus
(Drawing by E.S. Damstra,
published in Romero, 2003a).
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Fig. 2. Distributional map of the
Amblyopsid species (After Bergstrom, Nolti,
and Holtsford 1995) |
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Table 1.
Summary information on the amblyopsid species of fish.
Sources: Robison & Buchanan 1988, Page & Burr 1991,
Means & Johnson 1995, Pflieger 1997,
Romero 1998a,b,c, Romero & Bennis 1998.
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Species |
Maximum Size (Standard Length - SL, mm) |
Eyes |
Pigmentation |
Number
of Rays in Fins |
Pelvic
Fins |
Number
of rows of papillae in the caudal fin |
| Dorsal |
Anal |
Caudal |
| C. cornuta |
68 |
Microphthalmic |
Yes |
9-12 |
9-10 |
9-11* |
Absent |
0-2 |
| F. agassizii |
75 |
Microphthalmic |
Yes |
9-11 |
9-11 |
11-16* |
Absent |
0-2 |
| T. subterraneus |
75 |
Vestigial |
No |
7-10 |
7-10 |
10-15* |
Absent |
0-2 |
| A. spelaea |
110 |
Vestigial |
No |
9-11 |
8-11 |
11-13* |
Absent/very reduced |
4-6 |
| A. rosae |
65 |
Vestigial |
No |
7-9 |
8 |
9-11* |
Absent |
4-6 |
| S. poulsoni |
72 |
No vestiges? |
No |
9-10 |
8-9 |
21-22 |
Absent |
4 |
* Branched |
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