Sweepers

It's time to meet the sweepers.

Smallscale bullseyes Pempheris compressa, copyright John Turnbull.

Sweepers, Pempheridae, are a group of moderately sized marine fish (usually about fifteen to twenty centimetres in length) found around tropical reefs in the Indo-Pacific and western Atlantic. I don't know why they're called sweepers, but in some areas they may be among the most abundant fish on the reef. Distinctive features of the group include a short, high dorsal fin and a long anal fin. The lateral line is also distinctively long, extending past the end of the tail right onto the caudal fin. Perhaps the feature that most stands out about sweepers is their large eyes. The eyes are so big because sweepers are nocturnal; during the day they retreat into protected crevices and caves, emerging at night to feed on minute crustaceans and other small animals (Mooi 2001).

Pygmy sweeper Parapriacanthus ransonneti, from here.

Sweepers are divided between two quite distinct genera. Members of the genus Parapriacanthus have a more 'average fish-like' elongate profile with the body less deep than the head is long. The other genus, Pempheris, has a distinctively deep profile, deeper than the head is long. The exact number of species of pempherid appears to still be uncertain. Pempherids lack the striking markings of other tropical fish and species can appear very similar to each other. What is more, they have two layers of scales on the body, with the outer scales being larger than the inner and deciduous (easily shed), and loss of the outer scales has the potential to change an individual's superficial appearance. Early descriptions of pempherid species are often inadequate for their reliable identification, and new species continue to be described at a quite rapid pace. A recent publication by Randall & Victor (2015), for instance, described no less than thirty-four new species of Pempheris from various locations in the Indian Ocean, close to doubling the number of species in the genus at a stroke. The genus Parapriacanthus is much less diverse, with only about five recognised species.

Orange-striped bullseyes Pempheris ornata in hiding during the day, copyright Peter Southwood.

Because of their relatively small size and retiring habits, sweepers are mostly not that significant economically. At least one species, Pempheris xanthoptera, is fished off the coast of Japan and mostly eaten as fish paste; it is supposed to be quite tasty. Some have appeared in aquaria.

When foraging at night, sweepers communicate with each other by producing popping noises through muscular flexing of the swim bladder wall. Noise production increases in the presence of potential threats, perhaps to warn other members of the school. At least some pempherid species also have bioluminescent glands associated with the posterior part of the gut. The bioluminescent compound is not directly produced by the fish itself but obtained by consuming bioluminescent ostracods. I haven't found whether the function of this bioluminescence is specifically known for pempherids, but similar ventral glows in other fish provide camouflage by breaking up the fish's silhouette when seen from below.

REFERENCES

Mooi, R. D. 2001. Pempheridae. Sweepers (bullseyes). FAO Species Identification Guide for Fishery Purposes. The Living Resources of the Western Central Pacific vol. 5. Bony fishes part 3 (Menidae to Pomacentridae) pp. 3201–3204. Food and Agriculture Organization of the United Nations: Rome.

Randall, J. E., & B. C. Victor. 2015. Descriptions of thirty-four new species of the fish genus Pempheris (Perciformes: Pempheridae), with a key to the species of the western Indian Ocean. Journal of the Ocean Science Foundation 18: 77 pp.

Leucicorus: FAKE EYES!

In an earlier post, I told you about the fishes known as brotulas. These are one of the most prominent groups of fish in the deep sea. They tend not to be attractive fish: their lack of outstanding dorsal and tail fins makes them look like something between an eel and a cod, and like many deep-sea fishes they look somewhat flabby and lumpish. There are numerous genera of brotulas out there; the individual in the photo below represents the genus Leucicorus.

Leucicorus atlanticus, from Okeanos Explorer.

Leucicorus belongs to the brotula family Ophidiidae, commonly known as the egg-laying brotulas though Leucicorus' own reproduction has (so far as I have found) not been directly observed. The feature that most immediately sets Leucicorus apart from other brotulas is the eyes: Leucicorus species have very large eyes but the actual lenses are rudimentary or absent (Cohen & Nielsen 1978). It almost looks like they grew bigger and bigger to cope with the low light of the deep sea before they just kind of gave up at some point.

Two species of Leucicorus are currently recognised, each known from separate parts of the world. Leucicorus lusciosus is found in the eastern Pacific, whereas L. atlanticus is known from around the Caribbean. The two species differ in meristic characters and proportions: for instance, L. lusciosus has more dorsal and anal fin rays, but fewer vertebrae and gill rakers, and has a deeper body (Nielsen & Møller 2007). Leucicorus has also been found in the vicinity of the Solomon Islands, but interestingly enough Nielsen & Møller (2007) identified the specimen found as L. atlanticus rather than L. lusciosus, despite the latter species' more proximate distribution. One wonders if perhaps a third species is involved, yet to be recognised.

REFERENCES

Cohen, D. M., & J. G. Nielsen. 1978. Guide to identification of genera of the fish order Ophidiiformes with a tentative classification of the order. NOAA Technical Report NMFS Circular 417.

Nielsen, J. G., & P. R. Møller. 2007. New and rare deep-sea ophidiiform fishes from the Solomon Sea caught by the Danish Galathea 3 Expedition. Steenstrupia 30 (1): 21–46.

The Violet Demoiselle

Meet the violet demoiselle Neopomacentrus violascens (shown above in a photo by J. E. Randall). This little fish (adults get up to about 7.5 cm in length) is found in tropical waters of the western Pacific, from Japan in the north, south and east to northern Australia and Vanuatu. They usually associate in large schools around inshore reefs, and can commonly be found hanging around outcropping structures over soft bottoms such as coral or rocky outcrops, or wharf pilings (Koh et al. 1997). Violet demoiselles feed on small plant or animal plankton, such as copepods or algae.

The genus this species belongs to, Neopomacentrus, is one of the more recently recognised genera of the damselfish family Pomacentridae. It is similar to two larger genera in the family, Abudefduf and Pomacentrus, but differs from the former in having the hind margin of the preopercle (the anterior one of the bones making up the operculum or gill cover) crenulate or serrate rather than smooth. Pomacentrus has a similar preopercle, but has the suborbital region at least partially naked whereas Neopomacentrus has that region entirely scaly. Neopomacentrus violascens has a distinctive colour pattern, which is mostly a purplish brown, with bright yellow on the caudal fin and the rear of the dorsal fin.

According to Fishbase, individuals of this species form pairs when mating, and the females lay eggs that sink to the bottom and stick to the substrate. The eggs are then guarded and aerated by the males. I have come across reference to this species having been bred in captivity though I get the impression that they are not one of the most commonly kept aquarium fish. This may be because they are somewhat dull in coloration compared to related species, and they are fairly retiring in character. A recent blog post at Zoo Volunteer noted that damselfish species are rarely bred commercially due to the difficulty of providing suitable conditions. Instead, the market for species of this family is usually supplied with wild-caught individuals, commonly collected through the use of cyanide to essentially suffocate the fish until they lose conciousness. Not particularly pleasant for the fish, and arguably not that pleasant for the aquarist either as fish obtained in this method tend to have a much reduced lifespan.

REFERENCES

Koh, J. R., J. G. Myoung & Y. U. Kim. 1997. Morphological study on the fishes of the family Pomacentridae. I. A taxonomical revision of the family Pomacentridae (Pisces; Perciformes) from Korea. Korean Journal of Systematic Zoology 13 (2): 173–192.

Fishes be Crazy

Despite how it may look, there is absolutely nothing wrong with this fish. Butis butis is a moderately sized species of fish (up to about 14 cm in length) widespread in warmer waters around the Indian and western Pacific Oceans. They are found in shallow bays and mangrove swamps, often entering into estuaries and lower reaches of rivers. It is known by a range of vernacular names, including crimson-tipped gudgeon or duckbill sleeper. In the aquarium trade, it often goes by the name of crazy fish, in reference to its distinctive habit of swimming hanging vertically head-down or even swimming upside-down. Its predilection for such unusual angles assists it in remaining concealed from both predators and prey; Ryan (1981) noted that the addition of anaesthetic for collection purposes to a pool resulted in the sudden appearance of several specimens of which no sign had been previously seen. Their camouflage abilities are further enhanced by the ability to change colour to a certain degree, from pale to dark. Butis butis are ambush predators of smaller fish and invertebrates that they engulf in their broad jaws with rapid lunges.

Drawing of Butis butis, coloured from an original in Herre (1927) by M. L. Nievera.

The genus Butis belongs among the gobies, a somewhat notorious group of fish from a taxonomic perspective. In the words of the South African ichthyologist J. L. B. Smith, "The Gobioid fishes are one of the major trials of ichthyologists, and when general regional collections are worked up, these fishes tend to be pushed aside, and are apparently often identified with some impatience by those not specially interested" (Smith 1958). This notoriety is mostly due to the small size of many gobies, together with a tendency to the reduction of diagnostic features. Earlier authors classified Butis within the Eleotridae, a group of gobies (commonly known as sleepers, presumably due to their benthic habits) distinguished by having the pelvic fins separate from each other (in other gobies, the pelvic fins are united into a ventral sucker or disc). This, however, is a primitive feature only, and more recent molecular phylogenies have confirmed the paraphyly of Eleotridae in the broad sense. As a result, Butis and some of its nearest and dearest have been separated out into a separate family Butidae (Thacker 2011). However, while the separation of Eleotridae and Butidae seems to be fairly widely accepted, the two groups are still not clearly defined morphologically. Characteristic features of Butis relative to other gobies include (among others) the presence of a complete covering of scales, and a bony ridge above each eye (Smith 1958). In a number of species of the genus, including B. butis, the head is low and long, and the lower jaw distinctly protruding; however, the mudsleeper B. koilomatodon has a shorter, rounder head (this latter species, though originally native to a similar range to B. butis, has become invasive in more recent years in west Africa and Brazil, presumably carried in ballast water). Butis butis differs from other species in the genus in having small secondary scales at the base of most scales on the trunk, and the bony ridges above its eyes are more or less smooth (Herre 1927).

REFERENCES

Herre, A. W. 1927. Gobies of the Philippines and the China Sea. Philippine Bureau of Science Monographic Publications on Fishes 23: 1–352, 26 pls.

Ryan, P. A. 1981. Records of three new freshwater fishes from the Fiji Islands. Pacific Science 35 (1): 93–95.

Smith, J. L. B. 1958. The fishes of the family Eleotridae in the western Indian Ocean. Ichthyological Bulletin 11: 137–163.

Thacker, C. 2011. Systematics of Butidae and Eleotridae. In: Patzner, R. A., J. L. Van Tassell, M. Kovačić & B. G. Kapoor (ed.) The Biology of Gobies pp. 79–85. CRC Press.

Sea Bass, Mutant or Otherwise

...though to the best of my knowledge, none of them have fricking lasers on their heads.

Painted comber Serranus scriba, copyright Roberto Pillon.

The Serranidae are a group of marine fish that go by vernacular names such as sea bass, rock bass or rock cod. They are carnivores, and are found mostly around reefs in tropical and subtropical waters around the world. In appearance, they are fairly generalised (these are fish that look like fish) with a body shape that is longer than high, but not too long, and relatively big jaws with the lower jaw often jutting forward a bit beyond the upper. Some of them are quite colourful (as befits a tropical reef fish) and some of the smaller ones turn up in marine aquaria as a result. As used in the past, the Serranidae has been quite a broad grouping of fish united by having three spines on the margin of the opercle (the gill cover) and the maxilla in the upper jaw not hidden by the cheekbone when the mouth is closed. Members of this broad Serranidae were commonly divided between three subfamilies: the Serraninae (including the sea basses), Epinephelinae (including the groupers) and Anthiinae (basslets and goldies), though some authors further subdivided the Epinephelinae. However, recent molecular studies have indicated the polyphyly of this grouping, with the Serraninae and Epinephelinae occupying distinct positions within the clade known as the Serraniformes or Perciformes sensu stricto (see this old post), and so have cut the latter out of the Serranidae. As for the Anthiinae, their position remains uncertain, with some analyses placing them with the Serraninae and others with the Epinephelinae (Lautredou et al. 2013). As a result, a monophyletic Serranidae is probably to be restricted to the old 'Serraninae'.

Shy hamlet Hypoplectrus guttavarius, copyright Florent Charpin.

There are over eighty species listed for this restricted Serranidae on FishBase, but new ones continue to be described. As is common among reef fishes, it can be hard to determine exactly what counts as a species (whatever your preferred definition). A prime example of this is the genus Hypoplectrus, small serranids known as hamlets (no, I don't know why either) found in the Caribbean and the Gulf of Mexico. Hamlets come in a range of different colours and patterns, but structurally speaking the various forms are otherwise indistinguishable. As a result, some authors have regarded them as all colour morphs of a single species. Others have recognised close to twenty different species. Domeier (1994), conducting field observations on hamlets together with breeding experiments in the laboratory, found that different colour morphs would usually only mate with partners sharing their own colour pattern, though hybrid matings could be produced if no more suitable mate was provided. These hybrid matings produced offspring bearing intermediate colour patterns, and the rarity of such intermediates in the field led Domeier to infer that the different morphs were mostly acting as good species.

Kelp bass Paralabrax clathratus, photographed by Steve Lonhart.

Most sea basses are simultaneous hermaphrodites: they have both male and female reproductive organs functional at the same time. Though they are capable of fertilising their own eggs, they still usually breed in pairs with each individual alternating the release of male and female gametes. Not all serranids follow this reproductive template: members of the genera Chelidoperca and Centropristis are protogynous, starting their mature lives as females before switching over to males. Two species of Serranus, the lantern bass Serranus baldwini and the barred serrano S. psittacinus, are mostly simultaneous hermaphrodites like other species in the genus, but the largest individuals resorb their female organs and become exclusively males. Finally, many species of the genus Paralabrax have entirely separate males and females. Phylogenetic analysis suggests that protogyny may be the original mode of sexual development in the serranids, with separate sublineages developing simultaneous hermaphroditism vs separate sexes (Erisman & Hastings 2011). In correlation with this, individuals of Paralabrax that are functionally single-sexed have been found to retain non-functional remnants of the other sex's organs.

REFERENCES

Domeier, M. L. 1994. Speciation in the serranid fish Hypoplectrus. Bulletin of Marine Science 54 (1): 103–141.

Erisman, B. E., & P. A. Hastings. 2011. Evolutionary transitions in the sexual patterns of fishes: insights from a phylogenetic analysis of the seabasses (Teleostei: Serranidae). Copeia 2011 (3): 357-364.

Lautredou, A.-C., H. Motomura, C. Gallut, C. Ozouf-Costaz, C. Cruaud, G. Lecointre & A. Dettai. 2013. New nuclear markers and exploration of the relationships among Serraniformes (Acanthomorpha, Teleostei): the importance of working at multiple scales. Molecular Phylogenetics and Evolution 67: 140–155.

The Sculpins of Baikal

Drawing of Leocottus kesslerii, one of the more plesiomorphic of Baikal's sculpins, from here.

In a post that appeared on this site some seven years ago, I briefly introduced you to the sculpins of Lake Baikal. Sculpins, to quickly recap, are a group of bottom-dwelling fish found in Eurasia and North America, both in marine and freshwater habitats. At some point, a representative of the freshwater sculpins entered the massive Siberian lake known as Baikal, where it gave rise to one of the world's classic adaptive radiations.

To date, about thirty species of sculpin have been described from Lake Baikal. The level of morphological divergence between these species is such that they have been classified in the past into three separate families: while some were placed in the widespread family Cottidae, others were placed in two families endemic to Baikal, the Abyssocottidae and Comephoridae. However, phylogenetic analyses indicate that all the Baikalian sculpins originated from a single ancestor, and the entire clade is nested not only within the Cottidae but also within the genus Cottus (Kontula et al. 2003). Some of the Baikalian sculpins, such as the relatively basal Leocottus kessleri, retain a habitus and lifestyle similar to those of other sculpins elsewhere. Others, such as the golomyankas of the genus Comephorus, have become remarkably modified.

Specimens of Abyssocottus korotneffi, copyright Muséum National d'Histoire Naturelle.

The greatest diversity of Baikalian sculpins has resulted from their radiation into the lake's deep waters, which reach over 1600 metres (Sideleva 1996). This is a habitat unparalleled in any other freshwater lake. The only other great lakes reaching even comparable depths are the rift lakes Malawi and Tanganyika in Africa (the great lakes of North America, in contrast, are reasonably shallow). In the African lakes, the water quickly becomes anoxic below a fairly shallow top layer, and so the depths are devoid of multicellular life. Baikal, in contrast, is oxygenated all the way down (in this post, I speculated that this was due to Baikal's hydrothermal vents; it seems I was wrong. Baikal is oxygenated because the change in surface water temperature between summer and winter results in water circulating between layers and drawing oxygen down; in the tropical great lakes, where surface temperature remains fairly constant all year round, this circulation doesn't happen). The bulk of Baikal's deep-water sculpins make up the prior family Abyssocottidae, and exhibit adaptations similar to those seen in many marine deep-water fish. Their retinal structure has become simplified as a result of low light conditions. Their scales are reduced, and the lateral line system is composed of neuromasts exposed directly on the surface of the skin rather than contained in sub-surface canals and exposed to the outside environment via pores. The convergences between 'abyssocottids' and marine deep-sea fishes are so marked that some authors previously used them to argue for a direct marine ancestry of the Baikal fish (perhaps through a direct connection between Baikal and the sea that was once thought to have existed in the past), but this has been firmly quashed by the more recent molecular analyses. Instead, the majority of Baikal's deep-waters sculpins form a single clade that originated from shallower-water ancestors; the only exception is the genus Procottus, which includes both shallow-water and deep-water species (Kontula et al. 2003).

Golomyanka Comephorus dybowskii, from here.

Possibly sister to this deep-water clade are the aforementioned two species of golomyanka in the genus Comephorus. The golomyankas are without question the most bizarre members of the Baikalian sculpin radiation. They have become adapted to a pelagic mode of life, swimming in the open water column and feeding on Baikal's similarly remarkable pelagic amphipod Macrohectopus branickii (and as remarkable as Lake Baikal's sculpins are, they are nothing compared to its amphipods). It is not a simple matter for a sculpin to swim freely: they lost their swim bladders at an earlier stage in their evolution, so their native position is quite closely associated to the water's bed. To correct for this ancestral lack of buoyancy, golomyankas have lost their covering of scales and developed a low-density body structure that contains a high proportion of oil, about one-third of their total mass. Their pectoral fins have become greatly enlarged, covering about twice the area of the remainder of the body. The end result is that golomyankas are close to neutral buoyancy, and able to simply float in water column, waiting to ambush passing prey.

Golomyankas are also distinctive in their reproductive biology. Other sculpins lay their eggs in nests among stones, where they are tended by the male until they hatch. This includes the Baikalian genus Cottocomephorus, which has adopted a partially pelagic life comparable to that of Comephorus, but not to the same extent (Cottocomephorus species resemble Comephorus in having enlarged pectoral fins, but are otherwise more typically sculpin-like). Golomyankas, in contrast, are viviparous, releasing active larvae directly into the water column. Golomyankas are by far the most abundant fish in Lake Baikal, and a major component in the diet of other fish species (including, when young, other golomyankas). They are one of the key components in making Lake Baikal what it is, the world's only freshwater sea.

REFERENCES

Kontula, T., S. V. Kirilchik & R. Väinölä. 2003. Endemic diversification of the monophyletic cottoid fish species flock in Lake Baikal explored with mtDNA sequencing. Molecular Phylogenetics and Evolution 27 (1): 143–155.

Sideleva, V. G. 1996. Comparative character of the deep-water and inshore cottoid fishes endemic to Lake Baikal. Journal of Fish Biology 49 (Suppl. A): 192–206.

In a Pufferfish's Garden

Bullseye puffer Sphoeroides annulatus, copyright Geoffrey W. Schultz.

I don't know if it applies in other parts of the world, but one animal that you are guaranteed to see in the estuary here in Perth is pufferfish. One of the most instantly recognisable fish families, pufferfish (Tetraodontidae) are of course famed for their high toxicity, the determination of some people to eat them despite aforementioned toxicity, and their habit of swallowing air or water when threatened to inflate their distendible bellies. That last feature makes them a favourite of children (or at least of yours truly as a child), because their slow swimming style makes them one of the few fish that can be easily captured by hand (you just have to make sure you don't allow the fish to give you a nasty bite with their beak). The first feature makes them a lot less popular with fishermen who have to experience the frustration of reeling in a line to find that the bait has been taken by a puffer, then trying to remove the puffer from the hook while avoiding the aforementioned beak.

Oceanic puffer Lagocephalus lagocephalus, from Baino96.

There are a little under 200 known pufferfish species worldwide. Most of them are found in coastal marine and brackish waters, but there are also several species found in fresh water in South America, Africa and southeast Asia. Some marine species are also resistant to fresh water and may spend extended periods away from the sea. Some southeast Asian brackish-water Tetraodon species even make regular appearances in the the aquarium trade labelled as 'freshwater' puffers (Yamanoue et al. 2011), though their long-term survival requires more appropriate water conditions. The toxin associated with pufferfishes is not produced by the fish itself, but accumulated through its diet. As such, the exact level of toxicity of a pufferfish may vary according to season.

Grass puffer Takifugu niphobles, copyright OpenCage.

A molecular phylogenetic analysis of pufferfish by Yamanoue et al. (2011) identified four main clades in the family. These clades were also supported by a subsequent analysis by Santini et al. (2013), though the deeper relationships between the clades differed between the analyses. Yamanoue et al. (2011) identified a small number of freshwater clades (only one for each continent with freshwater taxa) and inferred that the transition from marine to fresh water had happened only rarely. Santini et al. (2013), in contrast, supported a higher number of transitions in tetraodontid history, though at least some of the difference between the two studies can be explained by differing definitions of 'freshwater'. For instance, some species of Takifugu usually live in brackish water but spawn in fresh water; Santini et al. counted these as freshwater species, but Yamanoue et al. did not.

Papuan toby Canthigaster papua, photographed by Dwayne Meadows.

One of the major clades identified within the Tetraodontidae includes the genus Lagocephalus, a group of relatively long-bodied puffers including some of the few pelagic puffer species. This genus may be the sister taxon of the remaining puffers (as found by Yamanoue et al.), or it may have a more nested position as sister to a clade including the mostly West Atlantic-East Pacific genera Sphoeroides and Colomesus (as found by Santini et al.). This latter clade includes South America's only freshwater puffer, the Amazon species Colomesus asellus. Santini et al. identified the basalmost tetraodontid clade as an Indo-West Pacific assemblage including the genus Takifugu and related taxa, which Yamanoue et al. had found as sister to the final clade including taxa related to the genus Tetraodon. This last clade includes the African and southeast Asian freshwater puffers (except for a few members of the Takifugu clade that cross into fresh water at times). It also includes the genus Canthigaster, the sharpnose pufferfish. In contrast to the more or less globular form of all other puffers, sharpnose puffers have a laterally compressed body form that superficially looks a bit more like a triggerfish than a puffer. Most Canthigaster species are reef-dwellers, a somewhat unusual habitat for a puffer (the other main group of reef-dwelling puffers being the genus Arothron, also in the Tetraodon clade).

Circular underwater 'nest' constructed by a pufferfish, from Spoon & Tamago.

One of the most remarkable characteristics of any puffer, though, was not discovered until quite recently. In 2012, it was announced that large structures observed off the coast of Japan by underwater photographer Yoji Ookata were in fact the work of pufferfish. These structures, circular and regular geometric patterns in the sea bed about 1.5 metres in diameter, were made by male puffers swimming against the sand. The structures are believed to function in attracting females, and also function as nests in which the females lay their eggs. Rather frustratingly, I haven't found any indication exactly which species of puffer is involved!

Puffer in the process of building a nest, also from Spoon & Tamago.

REFERENCES

Santini, F., M. T. T. Nguyen, L. Sorenson, T. B. Waltzek, J. W. Lynch Alfaro, J. M. Eastman & M. E. Alfaro. 2013. Do habitat shifts drive diversification in teleost fishes? An example from the pufferfishes (Tetraodontidae). Journal of Evolutionary Biology. doi: 10.1111/jeb.12112.

Yamanoue, Y., M. Miya, H. Doi, K. Mabuchi, H. Sakai & M. Nishida. 2011. Multiple invasions into freshwater by pufferfishes (Teleostei: Tetraodontidae): a mitogenomic perspective. PLoS ONE 6 (2): e17410. doi:10.1371/journal.pone.0017410.