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Home , Amblycipitidae, Anchariidae, Ariidae, Aspredinidae, Aspredo, Bagridae

American Fisheries Society Symposium 77:15–37, 2011 © 2011 by the American Fisheries Society

Global Catfi sh

JONATHAN W. ARMBRUSTER* Department of Biological Sciences, Auburn University 331 Funchess, Auburn University, Alabama 36849, USA

Abstract.—Catfi shes are a broadly distributed of freshwater fi shes with 3,407 cur- rently valid . In this paper, I review the different of catfi shes, all catfi sh fami- lies, and provide information on some of the more interesting aspects of catfi sh biology that express the great diversity that is present in the order. I also discuss the results of the widely successful All Catfi sh Species Inventory Project.

Introduction proximately 10.8% of all fi shes and 5.5% of all ver- tebrates are catfi shes. Renowned herpetologist and ecologist Archie Carr’s But would every one be able to identify the 1941 parody of dichotomous keys, A Subjective Key loricariid catfi sh pectegenitor as a to the of Alachua County, , begins catfi sh (Figure 2A)? It does not have scales, but it with “Any damn fool knows a catfi sh.” Carr is right does have bony plates. It is very fl at, and its mouth but only in part. Catfi shes (the Siluriformes) occur has long jaws but could not be called large. There is on every continent (even are known from a , but you might not recognize it as one as it Antarctica; Figure 1); and the order is extremely is just a small extension of the lip. There are spines well supported by numerous complex synapomor- at the front of the dorsal and pectoral fi ns, but they phies (shared, derived characteristics; Fink and are not sharp like in the typical catfi sh. How about Fink 1981, 1996). They are found in tropical and Pygidianops and Typhlobelus (Figure 3)? Pygidi- temperate freshwaters primarily, but some species anops are vaguely identifi able as catfi sh (although of predominantly freshwater families get into salt- they looks more like the leptocephalus larvae of water, and two families are predominantly marine elopiforms). However, Typhlobelus are very differ- (Burgess 1989). Across this broad range of , ent; they lack paired fi ns, so they obviously do not there is something that nearly everyone would be have spines, and even lack eyes. Specimens are only able to identify as a catfi sh. These fi shes are gener- about 30 mm long and a couple of millimeters in di- ally roughly cylindrical with a gray or silver body, ameter. There are no scales and the barbels are large, a large mouth, spines in front of the dorsal and or but can catfi sh be so small and featureless? pectoral fi ns, no scales, and, of course, the charac- With greater than 3,000 species (Table 1), per- ter that gives catfi shes their name, barbels (Fink and haps it should be expected that the diversity of cat- Fink 1981). fi shes is great, but this diversity is bewildering when Indeed, throughout the world, the catfi shes to we have such a stereotypical view of what it is to which people are most familiar fi t this general de- be a catfi sh (Arratia et al. 2003). It was with this in scription. In are the , mind that Larry Page, John Lundberg, Carl Ferraris, in and are the , in Europe Mark Sabaj, John Friel, and I developed the All Cat- and Asia are the , and in Australia, Papua fi sh Species Inventory (ACSI), a project ostensibly , and tropical to temperate continental to describe all catfi sh species. ACSI was funded shelves around the world are the . Because through a then new program at the U.S. National nearly every country on the planet has catfi sh, the Science Foundation called Planetary Biodiversity concept of catfi sh is well known to most people. But Inventories (PBIs) in 2003 and funding for ACSI this perception on the identity of catfi sh is limited ended in 2009. The PBIs are experimental programs because there are 37 families of catfi shes currently to accelerate . As species across the globe recognized and about 3,407 species (Table 1). Ap- disappear due to environmental degradation, it is * Corresponding author: [email protected] important to determine what species are out there. 15 16 ARMBRUSTER

FIGURE 1. Approximate global distribution of catfi shes based on Berra (2001), Ferraris (1999), Kailola (1999), and the map constructed by M. H. Sabaj-Perez for the All Catfi sh Species Inventory Web page. Dark areas are freshwater distributions (exaggerated in southeast Pacifi c islands) and gray areas are marine distribution (exag- gerated, generally limited to continental shelves). Extralimital records (crosses) from Grande and Eastman (1986), Lundberg (1975, 1992), Otero and Gayet (2001) and A. D. de la Peña (personal communication). The fossil record of catfi shes is reviewed in Gayet and Meunier (2003).

Without such basic knowledge, we cannot tell where and are supported as such mainly by the derived pres- to allocate resources in protecting the environment. ence of odontodes (integumentary teeth). These teeth Taxonomy is largely done piecemeal, and the PBIs range from minute denticles on the plates of loricari- are an attempt to bring together large teams of scien- ids and callichthyids to the sharp spines on the heads tists to develop the tools to speed up the process. In of blood-sucking to the long structures found this paper, I will review the families of catfi shes pro- on many breeding male loricariids. Given that loricar- viding their diversity, range, and basic ecology, and I ioids are sister to all other catfi shes, it is not surpris- will discuss the results of ACSI. The diversity of cat- ing that none of them fi t our basic image of a catfi sh. fi shes will follow the recent phylogeny of Sullivan et Despite this, 38.5% of catfi shes (1,312 species) are al. (2006) with additions by Lundberg et al. (2007) loricarioids. Loricarioids are found only in the neo- (Figure 4), although other phylogenies for catfi shes tropics and temperate regions of South America and are available (Mo 1991; de Pinna 1998; Hardman are broken up into fi ve families: Nematogenyidae, 2005). Teugels (2003) reviews the characters that , , Scoloplacidae, As- defi ne catfi sh families and major groups, and Diogo trobepidae, and . (2003) reviews phylogenetic relationships. Nematogenyidae (Nematogenyid Catfi sh) Basic Catfi sh Taxonomy Nematogenyidae consists only of from the mountains of Chile. The Nematog- Catfi shes are now believed to be in two distinct sub- enyidae is only one of two monotypic families of orders, the Loricarioidei and the Siluroidei, with catfi shes (the other is the recently described Lacan- currently unplaced. tuniidae).

Loricarioidei Trichomycteridae (Pencil or Parasitic Catfi shes) The loricarioids have been recognized as a monophy- Trichomycterids range throughout much of South letic group for a long time (Bailey and Baskin 1976) America and southern Central America. Trichmo- GLOBAL BIODIVERSITY 17

TABLE 1. Number of catfi sh species described (nominal species), currently valid, and the number added in the past 10 years. From Eschmeyer and Fong (2010). Nominal Valid Past 10 Taxonomy Common name Range species species years Loricaroidei Astroblepidae Naked or South America, 56 54 0 climbing catfi shes Panama Callichthyidae Armored catfi shes South America, 270 199 26 Panama Loricariidae Suckermouth armored South America, 936 813 174 catfi shes Costa Rica, Panama Nematogenyidae None Chile 3 1 0 Scoloplacidae Spiny dwarf catfi shes South America 5 5 1 Trichomycteridae Pencil or parasitic South America, 281 240 70 catfi shes Costa Rica, Panama Total Loricarioidei 1,551 1,312 271 Diplomystidae Velvet catfi shes Chile and 12 6 0 Siluroidei Akysidae Stream catfi shes 66 57 23 Torrent catfi shes South Asia 37 33 8 Amphiliidae Loach catfi shes Africa 98 80 16 catfi shes Madagascar 6 6 4 Ariidae Sea catfi shes Tropical to warm 343 147 15 temperate and nearby freshwaters Banjo catfi shes South America 67 39 3 Auchenipteridae Driftwood catfi shes South America 161 105 11 and Panama Auchenoglanididae Giraffe catfi shes Africa 37 25 1 Austroglanididae None Southern Africa 3 3 0 Bagridae Naked or bagrid Asia and Africa 314 210 44 catfi shes Whale catfi shes South America 50 42 20 Chacidae Squarehead, Southeast Asia 6 3 0 frogmouth, or angler catfi shes Clariidae Airbreathing or Southeast Asia and 216 111 14 walking catfi shes Africa None Africa 93 66 3 Cranoglanididae Armorhead catfi shes and 7 5 2 Thorny or talking South America 144 88 15 catfi shes None South and Central 289 199 15 America Heteropneustidae Air-sac catfi shes South Asia 8 4 0 Sun catfi shes South Asia 8 4 0 18 ARMBRUSTER

TABLE 1. Continued. Nominal Valid Past 10 Taxonomy Common name Range species species years Bullhead catfi shes 137 51 4 (Canada to Guatemala) Lacantuniidae Chiapas catfi sh Mexico 1 1 1 Malapteruridae Electric catfi shes Africa 27 19 14 Squeaker or Africa 261 204 22 upside-down catfi shes Pangasiidae Shark catfi shes South Asia 52 28 4 Pimelodidae Long-whiskered South America and 187 107 23 catfi shes Panama Plotosidae Eeltail or tandan Indo-Pacifi c, 80 40 5 catfi shes Australia, New Guinea Bumblebee catfi shes South America 40 34 8 Schilbidae None Africa and 120 61 4 South Asia Siluridae Sheatfi shes Europe and Asia 169 97 19 Hill-stream catfi shes South Asia 264 218 87 Total Siluroidei 3,291 2,089 385 Total Siluriformes 4,854 3,407 656

mycterids generally are equipped with interoper- Callichthyidae (Armored Catfi shes) cles and opercles that support odontodes. They can Callichthyids are distributed in Neotropical South evert these odontodes for protection and to provide America and southern Central America, most in the friction when climbing waterfalls. The candirus are . Most Corydoras are small (<50 often described as the only parasitic mm), detritivorous catfi shes covered in two rows of (Gudger 1930) and are in two subfamilies. The bony plates. Many species occur in hypoxic water feed on mucus and scales of other where they breathe air with the help of modifi ed intes- fi shes, and the Vandeliinae feed on blood of other tines capable of gas exchange (Persaud et al. 2006). fi shes. Fernández and Schaefer (2009) found that This type of air breathing may have been what helped arose only once in the trichomycterids. lead to the very unusual breeding strategy of Corydo- Vandellines swim into the gills of other fi shes and ras. It had been considered unusual that spawning use a uniquely derived median upper jaw bone to Corydoras. would sit in a T position, with the male as pierce the dorsal or ventral aorta and rapidly fi ll the bar of the T and the female with her mouth at the themselves with blood (Spotte et al. 2001; Spotte vent of the male as the post of the T. After apparent 2002; Zuanon and Sazima 2004). The fi shes then spawning, the female would move off and lay one to drink the blood and drop out of the fi sh when they several eggs, but it was unknown how the eggs were get their fi ll. They are often collected engorged fertilized. Kohda et al. (1995) found that the mode of with blood in sandy streams (Figure 5). Although fertilization is for the female to ingest the milt, move there are not that many species of hematophagous it quickly through her gastrointestinal tract, and de- trichomycterids, legends of the dominate posit the milt from her cloaca along with the eggs. our thoughts on this group of fi shes because they have been known to mistake the human urethra for Scoloplacidae (Spiny Dwarf Catfi shes) fi sh gills (Gudger 1930; Vezhaventhan and Jeyara- man 2007; Zuanon and Sazima 2004). The result is The Scoloplacidae consists of a single genus of damaging to both fi sh and human. very small fi shes (<25 mm). Scoloplacids are su- GLOBAL CATFISH BIODIVERSITY 19

FIGURE 2. Members of the Loricariidae: A. Pseudancistrus pectegenitor, B. anthrax (close- up of cheek odontodes in E and F), C. sp. mouth (pharyngeal jaws in G, CB—ceratobranchial, EB—epibranchial, UPJ—upper pharyngeal jaw), and D. macropthalmus male with tentacles. Images by M. H. Sabaj (A–D) and J. W. Armbruster (E–G). perfi cially similar to aspredinids; however, they of climbing waterfalls (Nelson 2006) by using their possess odontodes like other loricarioids, particu- pelvic fi ns, which are used almost in a walking mo- larly well developed on a plate on the snout (Bailey tion. Armbruster (2004) contends that the loricariid and Baskin 1976; Schaefer et al. 1989; Schaefer (from the Guiana Shield of 1990). They are found in fl oodplain lakes and small and Guyana and from the of Ven- streams and have been hypothesized to have a re- ezuela) is an astroblepid, but Schaefer (2003) and spiratory stomach (Armbruster 1998). Schaefer and Provenzano (2008) take an opposing view. According to Nelson (2006), the Astroblepidae Astroblepidae (Naked Suckermouth or Climbing is the most species-rich monogeneric . Catfi shes) Loricariidae (Suckermouth Armored Catfi shes) Astroblepids have a suckermouth, but are not armor- plated. They live in torrential streams of the Andes The Loricariidae (suckermouth armored catfi shes) and feed on larvae. Astroblepids are capable is the largest family of catfi shes with more than 20 ARMBRUSTER

FIGURE 3. The trichomycterids Pygidianops cuao (top) and Typhlobelus guacamaya (bottom) from Venezuela. Photos by N. K. Lujan.

800 species (23.9% of catfi shes). They are found wide, spoon-shaped structures that they use like in Neotropical South America and southern Cen- adzes to remove chunks of wood (Schaefer and tral America. The range of size and diversity of Stewart 1993; Armbruster 2003). But do they di- these fi shes is astounding. Fishes range from 20 to gest it? Nelson et al. (1999) suggest that they do; nearly 700 mm (with nearly this entire size range however, the wood looks the same in the anterior seen in one of approximately 16 species, the part of the intestine as the posterior part. Recently, group of Armbruster 2004). Males of German (2009) and German and Bittong (2009) many species have hypertrophied odontodes over did not fi nd that loricariids could digest wood various parts of their bodies, and in some species, because they lacked the ability to harbor sym- females have them as well. biotic microorganisms, the gut passage rate was Among the most unusual loricariids are mem- much too fast to digest wood, and there was not bers of Ancistrus, where males have long, fl eshy the right enzyme activity to digest wood. So, the tentacles on the tips of their snouts. We have hy- fishes may be consuming the wood only for the pothesized that these tentacles are larval mimics microorganisms that are themselves eating the (Sabaj et al. 1999) and are used by males to trick wood. females into thinking that they have young in their Also unusual among loricariids are mem- nests; this is because Ancistrus males protect the bers of the tribe Loricariina (Figure 2C). These young until they absorb the yolk sac, and having extremely dorsoventrally fl attened fi shes usually young may signify that the male is a good parent. have small tentacles around their mouths and very Recently, I have discovered what looks like larval few teeth. They do, however, have large, molari- mimics among the hypertrophied odontodes that form pharyngeal teeth (Figure 2G). I have found are held on evertible cheek plates in Ancistrus, as only seeds in the stomachs of the specimens I have well as closely related genera ( and dissected (Armbruster 2004), but some species are Pseudolithoxus). These cheek larval mimics have known to also consume insect larvae (Thomas and a rounded base that is yellow in life and a dark- Rapp Py-Daniel 2008). Although insectivory is ened region that proceeds distally from this base common in fi shes, loricariids are generally consid- and forks near the end of the odontode (Figure ered algivores (technically, this is incorrect as they 2E–F). mainly scrape biofi lm and should be classifi ed as The only two instances of lignivory (wood ). Granivory in fi shes is more unusual eating) among fi shes occur in the Loricariidae. but can be seen in many South American fi shes, Members of and the cochli- including other catfi shes (particularly doradids, odon species group have teeth modifi ed into short, Goulding 1980). GLOBAL CATFISH BIODIVERSITY 21

FIGURE 4. Phylogeny of the Siluriformes from Sullivan et al. (2006) and as modifi ed for the Big Africa clade by Lundberg et al. (2007). 22 ARMBRUSTER

FIGURE 5. The Trichomycterid sanguinea from Guyana with belly distended with blood. The fi sh is hanging by means of odontodes (integumentary teeth) on its interopercles and opercle. Photos by L. S. de Souza. GLOBAL CATFISH BIODIVERSITY 23

Diplomystidae (Velvet Catfi shes) species can be relatively easily collected by remov- ing hollow logs from rivers and letting the fi shes Diplomystids were found by Sullivan et al. (2006) fall out. Others, such as Ageniosus, are generally to be sister to the Siluroidei, but they did not pro- found in open water. Specimens of the unusual vide a higher ranking for the family. Diplomystids auchenipterid Gelanoglanis are less than 30 mm are from Chile and Argentina and are the only liv- standard length (SL; Figure 6A), and thus far, each ing catfi shes to retain maxillary teeth, a trait that discovered population appears to be a different spe- causes some to believe that diplomystids are the cies. These rare, pink fi shes only come out at night most primitive catfi shes (Arratia 1987; de Pinna where they can be seen swimming through the mid- 1998). dle of the water column. Male auchenipterids, like Siluroidei guppies, have a modifi ed anal fi n that they use for internal fertilization (Meisner et al. 2000). All of the remaining catfi sh families are siluroids, but the relationships of the families are poorly Doradidae (Thorny or Talking Catfi shes) known. Sullivan et al. (2006) found little structure Doradids are found in tropical South America. Al- to the relationships of the families but noted some though most doradids have a generalized catfi sh important clades, including the groups mentioned form, they generally have at least a single row of below. Although most siluroids are readily recog- plates with large spikes running down their sides nized as catfi shes, several groups push the defi ni- (Figure 6B). Birindelli et al. (2009) described the tion of catfi sh quite far. variation of doradids and found it to Cetopsidae (Whale Catfi shes) be extreme, with some bladders having many thin diverticula of unknown function coming off of a This small, South American family consists of the generally heart-shaped chamber. The function of whale catfi shes. Cetopsids are generally small (<60 these diverticula is unknown. mm, but some reach ~30 cm). Some cetopsids feed on larger fi shes by biting chunks out of their fl esh Heteropneustidae + Clariidae and swimming away (Burgess 1989). This clade has long been recognized, with hetero- Aspredinidae + Auchenipteridae + Doradidae pneustids occasionally placed within the Clariidae. This clade of South American catfi shes includes Heteropneustidae (Air-Sac Catfi shes) some generalized catfi shes (like Trachelyopterus, Heteropneustidae is from Southeast Asia and has Auchenipteridae) but generally are stranger fi shes just four species. Heteropneustes has one of the with many species armor-plated. most powerful toxins in catfi shes (Wright 2009), as Aspredinidae (Banjo Catfi shes) well as paired outgrowths of the buccal cavity that they use to breathe air (Munshi 1962). Aspredinids are found in tropical South America. The skin of aspredinids is keratinized and is shed Clariidae (Air-Breathing or Walking Catfi shes) like in reptiles (Friel 2009). These bizarre fi shes The clariids include a diverse array of species (Figure 6C) are often found in leaf litter, although from Asia and Africa. Most species are like the some prefer high montane streams and some, like walking catfi sh batrachus that has been , are found in and marine waters. introduced throughout the world (including Flor- Females of some species hold their eggs on out- ida), but some have elongated or anguilliform growths of their skin called cotylephores, which bodies like Chanallabes (Figure 7). Species of may aid in gas exchange for the embryos (Wetzel Clarias are able to leave water and walk to other et al. 1997). bodies of water. Unusual among fi shes, clariids can breathe air with their gills because they have Auchenipteridae (Driftwood Catfi shes) bony stays within the gills that keep them from Auchenipterids are found in tropical South Amer- collapsing in air (Graham 1997), and species of ica up to the Tuira River of Panama. They are of- Chanallabes are known to actually feed on land ten found in crevices or hollow logs. Many of the (Van Wassenbergh et al. 2006). 24 ARMBRUSTER

FIGURE 6. A. The auchenipterid Gelanoglanis sp. (Photo by P. Petri), B. the doradids armbrusteri (Photo by M. H. Sabaj), and C. the aspredinid Ernstichthys sp. (photo by N. K. Lujan).

Anchariidae + Ariidae the widest-ranging family of catfi shes, the Ariidae or sea catfi shes. Like the ariids, they have few, very The anchariids and ariids have long been thought to large eggs, but it is unknown what they do with the be related, with the anchariids occasionally placed eggs (Ng et al. 2008). in the Ariidae. Ariidae (Sea Catfi shes) Anchariidae (Madagascar Catfi shes) Ariids (Figure 8) are found on continental shelves Anchariids are a small group of fi shes found in off of all continents except Antarctica (Betancur freshwaters of Madagascar, but they are sister to and Armbruster 2009; Kailola 1999). Ariids are GLOBAL CATFISH BIODIVERSITY 25

FIGURE 7. A group of anguilliform clariids, Chanallabes sp. (Photo by J. W. Armbruster).

FIGURE 8. Unusual Papua New Guinea ariids: A. Doiichthys novaeguineae, B. Cochlefelis spatula, C. Brustia- rius nox, D. Pachyula crassilabris (photos by R. Betancur-R.). 26 ARMBRUSTER mouth-brooding fi shes and have very few, very Akysidae (Stream Catfi shes) large eggs; eggs up to 13 mm in diameter have Akysids are cryptically colored, small catfi shes been reported (Coates 1988). They are almost al- (generally less than 50 mm SL) from Southeast ways generalized catfi shes, but in the Indo-Pacifi c, Asia that have tubercles all over their bodies and they have their greatest diversity in morphology. extremely small eyes. Superfi cially, they resemble Perhaps the greatest diversity is ironically found aspredinids. There have been many new akysids in freshwaters of New Guinea, where ariids have described in the past 15 years due to collection ef- speciated tremendously (Allen 1991). The most forts in new places. unusual species that my graduate student (R. Be- tancur) and I collected in Papua New Guinea is Amblycipitidae (Torrent Catfi shes) Doiichthys, a small ariid (<200 mm) with a very fl attened head (Figure 8A). The Amblycipitidae is a small family of South- east Asian catfi shes that occur in swift waters as Big Asia far north as . They are generally elongate, resembling some members of the Heptapteridae or Sullivan et al. (2006) describe the “Big Asia” Trichomycteridae. Sullivan et al. (2008) corrobo- clade for the most speciose groups in Asia: the rated Hardman’s (2005) suggestion that the aky- Horabagridae, Bagridae, Akysidae, Amblycipiti- sids are not monophyletic and suggest that either dae, Sisoridae, and (now placed in the one new family needs to be described or that the Sisoridae). , generally considered a bagrid, was amblycipitids and akysids need to be placed in the not found to be related to other bagrids. Sisoridae. Horabagridae (Sun Catfi shes) Sisoridae (Hill-Stream Catfi shes) Horabagrids have been placed with the bagrids and Sisoridae is a large family of fastwater fi shes from schilbids up until recently. The family contains just Southeast Asia (Figure 9B–C). Among the more three genera and six valid species. Superfi cially, interesting adaptations in the Big Asia clade is the are similar to or to the bagrid presence of an attaching structure made of folds . Both species of Horabagrus are consid- of epidermis on the thorax in some sisorids, with ered threatened (Anvar Ali et al. 2007; Prasad et also having the pelvic fi ns connect- al. 2008). ed to forming a sucking attachment similar to that Bagridae (Naked or Bagrid Catfi shes) in gobies (Figure 9C). Bagrid diversity is centered in tropical Southeast Plotosidae (Eeltail or Tandan Catfi shes) Asia; however, the type genus are from The plotosids of the Indo-Pacifi c are the other ma- Africa (Mo 1991). Bagrids are mostly generalized rine family of catfi shes but are actually more di- catfi shes (Figure 9A), and many of the bagrids are verse in freshwater as in Australia and Papua New similar in appearance to the North American Ictal- Guinea (Allen 1991; Ferraris 1999). Plotosids have uridae or the South American Pimelodidae. heads that look like typical catfi shes, but gener- Some bagrids have unusual reproductive be- ally the adipose fi n has rays and is connected to haviors. are known to feed its young with the caudal and anal fi ns, giving the common name mucous secretions. Bagrus meridionalis of eeltail catfi shes. (Figure 10A). from have biparental care of nests of often lives in compact schools on coral reefs when young, the female produces unfertilized eggs to feed they are juveniles; thus, at least in the Indo-Pacifi c, the young, and the parents will even bring inverte- catfi shes are even reef fi shes. brates to the nest to feed the young (LoVullo et al. 1992). will often lay their eggs in the nests Chacidae (Squarehead, Frogmouth, or Angler of kampango, and both the cichlids and the catfi sh Catfi shes) will mutually defend the nest (McKaye 1985). Even more complexly, a clariid, Bathyclarias nyasensis, The Chacidae of Southeast Asia is a small family is known to occasionally brood parasitize kampango of three species that are ambush predators with a nests where the bagrid feeds the clariid young to the very large mouth (Figure 10B). Superfi cially, they detriment of its own (Stauffer 2010). are similar (although fl atter) than the North Ameri- GLOBAL CATFISH BIODIVERSITY 27

FIGURE 9. Members of the Big Asia clade: A. the bagrid (photo by S. Ferdous) and the sisorids B. yarrelli (photo by M. H. Sabaj), and C. Pseudecheneis longipectoralis showing fused pelvic fi ns and thoracic attaching region (photos by H. H. Ng). can fl athead catfi sh Pylodictus olivaris. Chacid are Siluridae (Sheatfi shes) called angler catfi shes because they are known to use their maxillary barbels as lures to attract fi shes Silurids range through much of Europe and Asia. for food (Roberts 1982). They differ from the typical catfi sh morphology 28 ARMBRUSTER

FIGURE 10. A. The plotosid Tandanus sp., B. (photo by P. J. Unmack), the chacid Chaca chaca (photo by S. Ferdous), and C. the silurid macrocephalus (photo by H. H. Ng). in that they lack dorsal spines and an adipose fi n istence, but morphological studies suggested that they and have a very long anal fi n. The family contains were three separate clades (de Pinna 1998). Molecular the longest catfi sh, wels glanis, which can studies put these families into a single clade, but they reach 5 m (Kottelat and Freyhof 2007). Silurids do are still recognized as separate families, and a fourth push the defi nition of the typical catfi sh a little bit, might be necessary for , which does not but perhaps the most unusual of them are the small fi t comfortably into any family (Sullivan et al. 2006). fi shes of Kryptopterus or glass catfi shes. Kryptop- terus have transparent bodies and live in schools in Pseudopimelodidae (Bumblebee Catfi shes) midwater. Their clear bodies make them diffi cult to The pseudopimelodids contain small fi shes like spot in turbid water (Figure 10C). and considerably larger fi shes such as Cephahlosilurus, which reach 40 cm and look sim- Pseudopimelodidae + Pimelodidae + Heptapteridae ilar to the North American fl athead catfi sh. Pseu- These three Neotropical families had been recognized dopimelodids are found in slow to swift water but in a single family, Pimelodidae, for most of their ex- are generally not found at higher elevations. GLOBAL CATFISH BIODIVERSITY 29

Pimelodidae (Long-Whiskered Catfi shes) undergo basin-wide migrations within the Amazon. This process takes many years, with juveniles mov- Pimelodidae are typical catfi shes for the most part, ing upstream to eventually spawn in the Andes (see with some notable exceptions (Figure 11). Baras and Lalèyè 2003 for review). (shovelnose catfi sh; Figure 11B) are particularly fl attened and some contain a very potent toxin in Heptapteridae (Heptapterid Catfi shes) their pectoral fi ns, and have a very fl attened snout (Figure 11D). Some pimelo- Heptapterids are the generally smaller species of dids like and the three-family group, and it is within this family

FIGURE 11. Members of the Pimelodidae: A. Brachyplatystoma rousseauxii, B. Sorubim lima, C. ornatus, and D. Platystomatichthys sturio (photos by M. H. Sabaj). 30 ARMBRUSTER

that the fi shes play around most with the general nests in Lake Tanganyika. Male cichlids mouth- catfi sh morphology. Perhaps the most unusual brood the eggs along with their own young, but is , a genus of phreatic catfi sh. The the catfi sh hatch earlier. Upon absorption of their phreatic zone is the region below the water table yolk sacs, the catfi sh young eat the larval cich- that is saturated with water, and these fi shes can lids (Sato 1986). The upside-down catfi shes (Syn- be collected in hand-dug wells but not in deeper, odontis) are unusual fi shes popular in the aquar- artesian wells (Muriel-Cunha and de Pinna 2005). ium trade that can be usually found swimming Heptapterids have a large range throughout South upside-down at the surface of the water. They are America and as far north as Mexico. actually aligning themselves with the substrate and will usually swim right-side up when near the Pangasiidae (Shark Catfi shes) bottom (called a ventral substrate response). An The Pangasiidae of Asia contains the Mekong experiment was done that subjected the fi shes to catfi sh Pangasianodon gigas, but also contains microgravity to see if this changed their behavior, the common food fi sh of the genus Pangasius but they still ignored light and responded only to (the or tra). Pangasius have been introduced surfaces (Anken and Hilbig 2009). into the U.S. supermarket, but not without back- Malapteruridae (Electric Catfi shes) lash from fi sh farmers. The U.S. Congress passed a temporary law that only the native Ictaluridae Perhaps the most unusual family of the Big Africa could be labeled catfi sh in grocery stores, and the clade is the Malapteruridae or electric catfi shes battle between North American fi sh farmers and (Figure 12B). Malapterurids have their lateral imported pangasiids continues. Juvenile Paga- musculature modifi ed into an electrical organ that sianodon are common in the pet trade under the is strong enough to shock prey and, from personal name “iridescent shark” (striped catfi sh Pangas- experience, cause quite a bit of pain to humans. Al- ius hypopthalmus), named because their dark bod- though for much of its history the Malapteruridae ies have a blue iridescence to them when young had been recognized as a single species, the diver- and a generalized shark shape. sity of form within the species was recently found to represent about 14 species in two genera. Big Africa Amphillidae (Loach Catfi shes) Sullivan et al. (2006) recognize a clade consisting of the African families Mochokidae, Malapteruri- Amphiliids are fast-water fi shes whose dae, Amphiliidae, Schilbidae, Auchenoglanidae members are rather typical catfi shes. Some of the ,and Claroteidae. Lundberg et al. (2007) found the derived species develop bony plates and an elon- newly discovered and described Mexican family gate appearance, with specimens of one genus, Lacantuniidae (Rodiles-Hernández et al. 2005) in Belonoglanis, looking much like a stick. this clade as well, but this family occurs in Mex- ico. Schilbidae () (Schilbid Catfi shes) The Schilbidae is a large family of mostly open- Mochokidae (Squeaker or Upside-Down Catfi shes) water fi shes that are superfi cially similar to silu- Mochokidae is the largest family in the Big Africa rids, and unlike other families in the Big Africa clade and had been considered related to the South clade, they range into southern Asia. Paralia, American families Doradidae and Auchenipteri- African glass catfi sh, are transparent, just as in dae to which they are superfi cially similar (Mo the glass catfi shes of Siluridae (Kryptopterus). 1991; de Pinna 1998). Mochokids often have dra- Given that we do often defi ne catfi shes based on matic patterns of spots that make them attractive the presence of a dorsal-fi n spine, it is unusual fi shes. Some mochokids have evolved that several schilbids lack spines (not unique to into similar morphologies as the South American schilbids), and some completely lack the dorsal loricariids by having a ventral mouth, jaws used fi n (Ailia and Paralia). There is currently an argu- for scraping submerged surfaces, and a sucker- ment among taxonomists on whether to spell the like mouth (Vigliotta 2008; Figure 12C). In anoth- family name Schilbidae or Schilbeidae (based on er instance of brood parasitism, the cuckoo catfi sh the genus ), and both names are commonly multipunctatus spawns over used (Nelson 2006). GLOBAL CATFISH BIODIVERSITY 31

FIGURE 12. Members of the Big Africa clade: A. Lacantunia enigmatica (photo by J. W. Armbruster), B. Mala- pterurus monsembeensis, and C. asymetricaudalis (photos by J. P. Friel).

Auchenoglanididae (Giraffe Catfi shes) Claroteidae (Claroteid Catfi shes) Auchenoglanidids and claroteids are often placed Claroteids are the large catfi shes of Africa. They look in the same family, but the recognition of Lacantu- superfi cially similar to Bagrus and were once consid- niidae as the sister to the Claroteidae would make ered to be bagrids. Most species are in , this family paraphyletic. Auchenoglanidids are most which has its greatest diversity in the Congo. The similar in appearance to mochokids but typically most unusual claroteids are species of the Lake Tan- have a longer snout. ganyika endemic genus , the fi rst known fi shes to have bipaternal mouth brooding (Ochi et al. 2000). 32 ARMBRUSTER

Lacantuniidae (Chiapas Catfi sh) Cranoglanididae (Armorhead Catfi shes) Lacantuniidae was recently described for the new- (Figure 13A) has just three species ly discovered Lacantunia enigmatica of Chiapas, from China and Vietnam. The fi shes look similar to Mexico. Chiapas catfi sh L. eniqmatica sort of the North American bullheads of the genus Amei- looks like a large (Ictaluridae) but with urus. a different number of barbels (Figure 12A). The disjunct distribution of Lacantuniidae and its pro- Ictaluridae (Bullhead Catfi shes) posed sister family (Claroteidae) is unusual and, if Despite the diversity of catfi shes around the world, the relationship is true, requires explanation. The the waters of North America north of Mexico are split between Lancantunia and claroteids occurred relatively depauperate in catfi sh morphologies. All after the breakup of Gondwanaland; thus, relatives ictalurids (even the boldly patterned piebald mad- of Lancantunia and or Claroteids would have had tom Noturus gladiator, Figure 13B) look like typi- to have gone extinct over large parts of the globe cal catfi shes. As this volume can attest, the ictalurids (Lundberg et al. 2007). (particularly Ictalurus and Pylodictis) are probably the most widely studied catfi shes in the world. Also, Cranoglanididae + Ictaluridae given the fact that there are similar fi shes on every Last in Sullivan et al.’s (2006) analysis is the clade continent, the North American research will provide that includes the North American bullhead catfi sh- a lot of insight into the dynamics of catfi sh popula- es or Ictaluridae. Ictalurids are found to be related tions and fi sheries around the world. This insight is to the only genus of cranoglanidid, Cranoglanis of limited by the great diversity of form, , be- China and Vietnam. This relationship is interesting haviors, and ecology of catfi shes. because several groups of (such as paddle- Austroglanididae (Austroglanidid Catfi shes) fi shes , alligators, and many other groups) share a similar distribution. The linkage between eastern The three species and one genus of austroglani- North America and China is sometimes referred to dids are known from southern Africa. No austro- as a Grayan distribution after the botanist Asa Gray glanidids were available for either of the main who published on it in the 1800s (Gray, 1846; Pe- molecular studies of catfi shes (Hardman 2005; tersen and Hughes 2007). Sullivan et al. 2006), and morphological hypoth-

FIGURE 13. A. The cranoglanid Cranoglanis sinensis (Photo by H. H. Ng), and B. the ictalurid Noturus gladi- ator (Photo by M. A. Thomas). GLOBAL CATFISH BIODIVERSITY 33 eses differ as to their placement. Mo (1991) sug- the past 10 years (Table 1). ACSI has been active gests that they are related to cranoglanidids of for much of the 10 years, and we can see that the Asia, and de Pinna (1998) suggests that they are number of catfi sh species has increased by nearly sister to a complex, pan-tropical clade of hora- 24% in the past 10 years. bagrids, claroteids, schilbids, pangasiids, some We also developed many other products, in- bagrids, and pimelodids. cluding a database of around 10,000 photographs (mostly types) on our Web site, which enables peo- The Role of the All Catfi sh Species Inventory ple to see the types without having to ask for them on loan or to travel to the diverse places around the ACSI was conceived in the fi rst round of submis- world where types are located. We began assembling sions for NSF’s Planetary Biodiversity Inventory PDFs of published papers on the Web site as well program, was funded in 2003, and went on to be because a lot of the old literature is inaccessible to widely successful. Although the PBIs were fash- many people around the world. We developed aux- ioned with the idea of discovering and describing iliary sites on the diversity of some catfi sh families all species in a clade dispersed across the planet, it (www.fl mnh.ufl .edu/fi sh/freshwater/catfi sh/catfi sh_ is unreasonable to think that all undescribed spe- home.htm) as well as an online atlas of catfi sh bones cies of catfi shes could be described in the 6 years (http://catfi shbone.acnatsci.org). Through funds to in which ACSI was active. What the PBIs really participants, as well as funds in the laboratories of serve to do is to make taxonomy easier. Our re- the principal investigators, we funded fi eldwork in sults were dramatic, we developed a team of 460 Asia, Africa, the Australian continent (Papua New participants from 53 countries and together we Guinea), and especially South America, as well as have increased the rate of species descriptions, museum visits for researchers to complete taxonom- especially when compared with other otophysan ic revisions. Many of the participant awards went fi shes (Figure 14). Eschmeyer and Fong (2010) to graduate students, as the only way to assure the keep a list of all of the described and valid spe- completion of taxonomy is to develop the next gen- cies by family, as well as the species described in eration of taxonomists.

FIGURE 14. Percent change in rate of species description in the three largest orders of primary freshwater fi shes in the fi ve main years of All Catfi sh Species Inventory (2004–2008) compared to the previous 5 years (1999–2003). 34 ARMBRUSTER

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