Field of Science

Giant Centipedes (That Aren't All Giants)

Scolopendra morsitans, copyright Jiri Lochman/Lochman Transparencies.


It was a dark night, but not stormy (nights tend to be dark, as a rule). We were out collecting for our regular survey when we encountered a large centipede (the same species as in the photo above) crossing the road, and decided to add it to our collection. Taking out the large 20-cm forceps that we had on hand for dealing with venomous animals, one of us used them to grab the centipede.

The response was electric. Rather than trying to escape its attacker, the centipede instantly whipped back and lashed itself around the forceps, doing its best to bite into them. Had the actual wielder of the forceps been within its reach, they would have been in for a world of pain. When dealing with scolopendrid centipedes, you should always remember three things: they are big, they are fast, and they are mean.

Scolopendridae are unmistakeable. They include the giants of the centipede world, with the largest species (the South American Scolopendra gigantea) reaching up to a foot in length. Even the smaller species are relatively robust compared to other centipedes. Like all centipedes, the first pair of legs is modified into a robust pair of 'fangs' used for delivering venom (when I referred above to a centipede 'biting', this is what I was properly referring to). Large scolopendrids have the most dangerous centipede stings, potentially causing intense pain, though fatalities are very rare (Bush et al., 2001, noted that no centipede fatalities were known from the US, though they did refer to a single known child fatality in the Philippines). Their hunting prowess is amply demonstrated in this video of a large scolopendrid hunting bats by quite literally snatching them out of the air:
ARKive video - Amazonian giant centipede hunting bats inside a cave


But lest you think that scolopendrids are all venom and viciousness, let me point out that they also have their endearing qualities. Female scolopendrids make devoted mothers, coiling around their egg clutches and regularly cleaning them to prevent fungal attack. Even after the eggs hatch, the female continues to coddle and groom her young. Brunhuber (1970) recorded that females of Cormocephalus anceps spent at least three months (from late September to late December) caring for young before they struck out on their own. Even after becoming independent, the young do not reach sexual maturity until they are at least two years old. Other scolopendrids may mature more quickly, at about one year (Lewis 1972). Individual centipedes may live for several years.

Female Scolopendra morsitans cleaning her eggs, copyright H. J. B..


The Scolopendridae belong to a larger centipede group called the Scolopendromorpha. Most scolopendromorphs have bodies with 21 or 23 leg-bearing segments, except for one remarkable scolopendrid species from central Brazil that has 39 or 43 leg-bearing segments (Chagas-Junior et al. 2008). Non-scolopendrid species are often much smaller than the Scolopendridae, with some being only about 10 mm in length. These smaller scolopendromorphs also differ in eye morphology: Scolopendridae have a patch of four ocelli on either side of the head, but other scolopendromorphs are mostly blind and lack ocelli. In the past the blind scolopendromorphs have been treated as a single family Cryptopidae, but recent authors have mostly recognised three separate families Cryptopidae, Scolopocryptopidae and Plutoniumidae in light of uncertainty about the monophyly of a broader Cryptopidae. Nevertheless, a recent phylogenetic analysis by Vahtera et al. (2012) combining both morphological and molecular data did support a single blind clade. Unfortunately, no phylogenetic analysis to date has been able to include Mimops orientalis, an odd scolopendromorph known from a single specimen collected in China in 1903 and placed by Lewis (2006) in its own distinct family. Mimops possesses but a single ocellus on either side of the head, potentially making it very intriguing for the question of whether blindness has evolved in scolopendromorphs more than once.

The blind scolopendromorph Scolopocryptops sexspinosus, copyright Troy Bartlett.


REFERENCES

Brunhuber, B. S. 1970. Egg laying, maternal care and development of young in the scolopendromorph centipede, Cormocephalus anceps anceps Porat. Zoological Journal of the Linnean Society 49 (3): 225-234.

Bush, S. P., B. O. King, R. L. Norris & S. A. Stockwell. 2001. Centipede envenomation. Wilderness and Environmental Medicine 12 (2): 93-99.

Chagas-Junior, A., G. D. Edgecombe & A. Minelli. 2008. Variability in trunk segmentation in the centipede order Scolopendromorpha: a remarkable new species of Scolopendropsis Brandt (Chilopoda: Scolopendridae) from Brazil. Zootaxa 1888: 36-46.

Lewis, J. G. E. 1972. The life histories and distribution of the centipedes Rhysida nuda togoensis and Ethmostigmus trigonopodus (Scolopendromorpha: Scolopendridae) in Nigeria. Journal of Zoology 167 (4): 399-414.

Lewis, J. G. E. 2006. On the scolopendromorph centipede genus Mimops Kraepelin, 1903, with a description of a new family (Chilopoda: Scolopendromorpha). Journal of Natural History 40 (19-20): 1231-1239.

Vahtera, V., G. D. Edgecombe & G. Giribet. 2012. Evolution of blindness in scolopendromorph centipedes (Chilopoda: Scolopendromorpha): insight from an expanded sampling of molecular data. Cladistics 28: 4-20.

The World's Scorpion

Pair of lesser brown scorpions Isometrus maculatus in captivity, from here. The more elongate individual at lower left is a male; his stouter companion is female.


With their elongate, sting-tipped tails, scorpions are instantly distinguishable from any other arachnid. Depending on how you look at it, they are either charismatic or infamous (not many invertebrates have constellations named after them). Yet though the world diversity of scorpions is not unrespectable (about 1750 species have been described so far), distinguishing one scorpion species from another can be a challenging prospect. They tend, as a whole, to be a morphologically conservative group.

As a result, new species of scorpion continue to be described, at a rate limited only by the relatively small number of people taking up the challenge. Isometrus is a genus of about thirty species of scorpion found mostly from in southern Asia and Australasia, from Pakistan to northern Australia and New Caledonia. A good third of those species have only been described since 2000, and probably more remain to be described. They are small, relatively slender scorpions, often with the 'fingers' of the chelae noticeably darker than the 'palms' (as in the photo above). They have a sting that has been referred to as painful, but doesn't seem to have lead to recorded fatalities in humans.

Isometrus has been divided between two subgenera, Isometrus sensu stricto and Reddyanus (Kovařík 2003). The two subgenera are distinguished primarily by the locations of trichobothria, long sensory hairs, on the chelae. The higher diversity of species belong to Reddyanus, but Isometrus is by far the more widespread subgenus. This is due to its inclusion of one particular species: I. maculatus, commonly given the rather underwhelming name of 'lesser brown scorpion'. For some reason, Isometrus maculatus has proven itself very amenable to transport by humans. It may have been originally native to Sri Lanka (as cited here) but from there it has spread to tropical regions around the world. It is found in North America, South America, Africa and Australia, and on various oceanic islands such as Hawaii, Saint Helena and the Seychelles. In Europe, it is only known from southern Spain, though it may have been the species originally indicated by Linnaeus' 'Scorpio europaeus' (Fet et al. 2002; some authors have consequently used the name 'Isometrus europaeus' for this species, but Linnaeus' name was declared invalid by the ICZN due to the uncertainty of its identity). Currently, I. maculatus is regarded as the world's most widespread scorpion species. No other Isometrus species has been subject to the same degree of spread, though I am personally inclined to wonder about the distribution of I. heimi, recorded by Kovařík (2003) from both New Guinea and New Caledonia.

REFERENCES

Fet, V., M. E. Braunwalder & H. D. Cameron. 2002. Scorpions (Arachnida, Scorpiones) described by Linnaeus. Bull. Br. Arachnol. Soc. 12 (4): 176-182.

Kovařík, F. 2003. A review of the genus Isometrus Ehrenberg, 1828 (Scorpiones: Buthidae) with descriptions of four new species from Asia and Australia. Euscorpius 10: 1-19.

Lachenalia

Back in 2011, I presented you with a post on the southern African flowering bulb genus Ledebouria. In that post, I mentioned that Ledebouria was just one of a wide diversity of ornamental plants found in that part of the world.

Lachenalia elegans var. flava, from the Pacific Bulb Society.


Lachenalia, sometimes known as Cape cowslips, is a genus of over 100 species found in Namibia and South Africa. Most Lachenalia species sprout and flower in the winter. Lachenalia is not too distant a relative of Ledebouria—both are classified in the squill tribe Massonieae—and bears a distinct resemblance to the latter with its fleshy leaves that are often blotched with purple. Some species of Lachenalia share the geophyllous habit I described in the earlier post for some Ledebouria, with the leaves growing pressed closely to the ground. However, Lachenalia differs from Ledebouria in having flowers with well-developed bracts, and anthers arranged in two series. Also, while the scales of Ledebouria bulbs are often loose, though of Lachenalia bulbs are always tightly packed (Manning et al. 2004).

Lachenalia zebrina f. zebrina, photographed by Alan Horstmann.


Lachenalia species include some popular garden plants, to the extent that some are known as invasive weeds here in the Perth region. Nevertheless, a simple image search immediately shows why they are so popular. Varieties of this genus are available in reds, pinks, yellows, purples... One species, L. viridiflora, has flowers of a quite remarkable turqouise colour. Though revered in cultivation, L. viridiflora is critically endangered in the wild, with a range of only 19 km2 in which it is threatened by grazing, housing development and (almost ironically) the collection of specimens for horticulture.

Lachenalia viridiflora, photographed by A. Harrower.


REFERENCE

Manning, J. C., P. Goldblatt & M. F. Fay. 2004. A revised generic synopsis of Hyacinthaceae in sub-Saharan Africa, based on molecular evidence, including new combinations and the new tribe Pseudoprospereae. Edinburgh Journal of Botany 60 (3): 533-568.

There's No Such Thing as Caddids

Caddo agilis, from here.


Long-time readers of this site may recall my previous rants on the subject of the prolific, but not entirely reliable, arachnologist Carl-Friedrich Roewer. Hopefully, this post will serve to rehabilitate Roewer's image a little, because occasionally something comes along about which he was right in the first place.

Among Roewer's innovations in Die Weberknechte der Erde, his 1923 revision of the world Opiliones fauna, was the introduction of a new family for Acropsopilio, an odd little harvestman from South America. He placed this new family in the Dyspnoi, a subgroup of the Palpatores (long-legged harvestmen) that is otherwise found in Eurasia and North America. Acropsopilio was a distinctive beast, a tiny harvestman with relatively massive eyes (just take a look at the picture below!) Over time, other authors added to the Acropsopilionidae: species are now known from Australia, New Zealand and South Africa. They are nowhere comon, though.

Specimen of Acropsopilio neozelandiae, photographed by Stephen Thorpe.


In 1975, the acropsopilionids were revised by Shear (1975), who proposed that they were related to Caddo, a genus of harvestmen found in north-eastern Asian and north-eastern North America. That's not a typo, by the way: the range of this genus includes Japan and New England, but not the spaces in between. To make things just that extra bit wierder, the genus includes two species, C. agilis and C. pepperella, both of which are found in both the sections of its overall range. Genetic analysis has demonstrated that this wierdness is real, and not just convergence or one variable species (Shultz & Regier 2009). Caddo had previously been classed as a member of the Eupnoi, the other main subgroup within the Palpatores, but resembled acropsopilionids in features such as the small size and large eyemound. Shear proposed classing them all as a single family, Caddidae, with two subfamilies: one for Caddo and one for the Acropsopilioninae. Subsequent authors have followed his lead, and the Caddidae has come to be placed within the Eupnoi as the sister taxon to the Phalangioidea (the group including the familiar long-legged harvestmen such as the field harvestman Phalangium opilio).

Nevertheless, there was still a bit of humming and hawing going on behind the scenes. Despite the overall similarities in habitus between Caddo and acropsopilionines, several of the finer details (such as the structure of the pedipalps and genitalia) were quite different. Phylogenetic studies commenting on the position of caddoids within the Opiliones had generally included Caddo only, and not included any representatives of the acropsopilionines. And so it is quite welcome to see a new publication by Groh & Giribet (in press) in which they produced a molecular phylogenetic analysis of the caddids as a whole. The result, as hinted in the first paragraph, is that the caddids are not supported as a monophyletic group. Caddo remains in its accustomed position within the Eupnoi, but the acropsopilionids are placed as the sister clade to the Dyspnoi. Roewer, it turns out, had them in the right place to begin with.

This has some interesting implications: for instance, the otherwise entirely Holarctic Dyspnoi have just acquired a Gondwanan basal group. Also, the large eyemound is either a convergent feature between Caddo and acropsopilionines, or a retained primitive feature from the palpatorean common ancestor. Groh & Giribet suggest the latter, but I suspect the former to be just as likely (it may be related to small size: some phalangioids, such as the Mediterranean Platybuninae and the Western Australian Megalopsalis tanisphyros, also have large-ish eyemounds). But the greatest surprise for yours truly was something else: one particular 'acropsopilionine' genus, Hesperopilio, was not placed either with Caddo or the other acropsopilionines. Instead, it was placed closer to the the phalangioid family Neopilionidae: the subject of my own research.

When I produced my revision of the Australasian phalangioid family Monoscutidae (which I ended up synonymising with Neopilionidae), I included Caddo as an outgroup taxon in my morphological phylogenetic analysis. At the time, my supervisor asked me why I didn't include an acropsopilionine as well, but I demurred on two points. One was that, as rare as acropsopilionines were in collections, males were even rarer (there is evidence that they are commonly parthenogenetic, as for that matter is Caddo). The other was that acropsopilionine genitalia were truly bizarre, and I couldn't determine which parts of the acropsopilionine penis corresponded to where on the monoscutid organ.

I was basing that judgment on Acropsopilio and the South African genus Caddella (offhand, there is a longstanding tradition in harvestman taxonomy that whenever the name Caddella appears in a paper, it must be mis-spelled at least once). I still stand by that judgment. But upon seeing the results of Groh and Giribet's molecular analysis, I looked up the description of Hesperopilio (Shear 1996), which includes a drawing of the male genitalia. And suddenly, I was struck by the possibility that they could indeed be neopilionid-like. So I tried entering Hesperopilio into my original data set using the published descriptions. The result? Though missing a fair amount of data (my coding would need to be checked against actual specimens), a rough run suggests that morphology supports Hesperopilio as a neopilionid too!

The simplified version of what I end up with. Remember, this is by no means a thoroughly vetted result; this is just me going "what if I do this?"


So let that be a lesson, I suppose. Because of the belief that Hesperopilio was an acropsopilionine, I had never even considered taking a closer look at it. As it turns out, I really should have!

REFERENCES

Groh, S., & G. Giribet (in press) Polyphyly of Caddoidea, reinstatement of the family Acropsopilionidae in Dyspnoi, and a revised classification system of Palpatores (Arachnida, Opiliones). Cladistics.

Shear, W. A. 1975. The opilionid family Caddidae in North America, with notes on species from other regions (Opiliones, Palpatores, Caddoidea). Journal of Arachnology 2: 65–88.

Shear, W.A. 1996. Hesperopilio mainae, a new genus and species of harvestman from Western Australia (Opiliones: Caddidae: Acropsopilioninae). Records of the Western Australian Museum 17: 455–460.

Shultz, J. W., & J. C. Regier. 2009. Caddo agilis and C. pepperella (Opiliones, Caddidae) diverged phylogenetically before acquiring their disjunct, sympatric distributions in Japan and North America. Journal of Arachnology 37: 238–240.

Rock Mosses

Black rock-moss Andreaea rupestris, photographed by Sture Hermansson.


Upon first examining the picture above, you might think that you were looking at a patch of moss. Which would not be an unreasonable assumption to make, because that is exactly what you are looking at. But this is not just any moss, but perhaps one of the most interesting mosses out there.

The hundred or so species of the genus Andreaea, found in cooler regions around the world, are commonly known as rock mosses or granite mosses in reference to their preferred growth habitat on acidic rocks. They can often look black or red rather than green (presumably relative to how dry they are), and they are often brittle. Glime (2013) notes that a characteristic feature of granite mosses is that a hand brushed over one will come away with small fragments stuck to it, and suggests that this may act as a method of vegetative dispersal. Usually, granite mosses are autoicous: a single plant has both male and female reproductive structures, but they are borne in separate clusters. What makes Andreaea really interesting, though, is how it produces spores. As explained in the diagram I used in an earlier post, mosses produce spores from a sporophyte (diploid plant) that grows supported by the gametophyte (haploid plant) that comprises the green, vegetative stage of the moss life cycle. In most mosses, the sporophyte holds itself up by means of a long stalk called a seta, and spores are released from the terminal capsule by the ejection of a covering operculum.

Capsules of Andreaea, photographed by David Tng.


Andreaea does things differently. In this moss, the capsule is not supported by its own stalk, but is instead lifted up on an extension of the gametophyte called a pseudopodium. And instead of popping off an operculum, the Andreaea capsule splits open longitudinally into a squashed crown. Andreaea shows its difference after the spores are released as well: while the spores of other mosses germinate into a filamentous protonema (the moss 'seedling', as it were), the protonema of Andreaea bears thalloid appendages.

Such is the distinctive of Andreaea that it has been classified in a separate class from most other mosses, the Andreaeopsida. Phylogenetic analysis has demonstrated that Andreaea is one of the earliest diverging of all mosses, being the next to diverge from the main moss lineage after the Sphagnopsida (the group that includes Sphagnum). In some classifications, the class Andreaeopsida is restricted to Andreaea alone, but there are two other small genera that have been grouped with it in the past.

Andreaeobryum macrosporum, photographed by Masaki Shimamura.


Andreaeobryum macrosporum is a single unusual moss species found in north-west North America. Like Andreaea, it is found growing in rocks, though its preference is for basic rocks such as limestone. It also resembles Andreaea in having a spore capsule that opens through slits, though in the case of Andreaeobryum the apex of the capsule eventually wears off as well and the capsule splays fully open. The biggest difference between Andreaea and Andreaeobryum is that the capsule of the latter is not raised on a gametophytic pseudopodium, but possesses its own supporting seta like that of typical mosses (albeit a particularly short and stubby one). The phylogenetic position of Andreaeobryum remains uncertain: a molecular analysis by Chang & Graham (2011) recovered an Andreaea-Andreaeobryum clade, but with very low support.

Takakia lepidozioides, from here.


The real wild-card in basal moss phylogeny, however, is the little green monster known as Takakia. Takakia is a genus of two species found in western North America and eastern Asia. It was first discovered in the Himalayas and described in 1861—as a liverwort. This is a bit like being presented with a new species of snake, and describing it as a type of eel. But it has to be pointed out that Takakia possesses some very un-moss-like features. It has finely divided leaves, a feature common in liverworts but not known from any other moss. Its leaves contain oil bodies: again, unlike any other moss, but like many liverworts. Matters were not helped by the fact that Takakia was first described from vegetative material only, and it was not until the description of sporophytes in the 1990s that Takakia was conclusively accepted as a moss (Renzaglia et al. 1997).

Even so, its position within the mosses remained uncertain. A relationship with Andreaea has been suggested, as the capsule (though borne on a seta rather than a pseudopodium) opens through a single spiral slit. However, recent phylogenetic analyses have not supported a direct relationship between the two. Chang & Graham (2011) found in the analysis of their data that its position was vulnerable to the analytical model used: it could be placed as the sister taxon to all other mosses, or it could be the sister to the Sphagnopsida (with the two together being the basalmost moss clade). We have not heard the last of little Takakia.

REFERENCES

Chang, Y., & S. W. Graham. 2011. Inferring the higher order phylogeny of mosses (Bryophyta) and relatives using a large, multigene plastid data set. American Journal of Botany 98 (5): 839-849.

Glime, J. M. 2013. Bryophyta - Andreaeopsida, Andreaeobryopsida, Polytrichopsida. Chapt. 2-6. In: Glime, J. M. Bryophyte Ecology. Volume 1. Physiological Ecology. Ebook sponsored by Michigan Technological University and the International Association of Bryologists. Last updated 29 June 2013 and available at .

Renzaglia, K. S., K. D. McFarland & D. K. Smith. 1997. Anatomy and ultrastructure of the sporophyte of Takakia ceratophylla (Bryophyta). American Journal of Botany 84 (10): 1337-1350.

The Diversity of Neosauropods, or Pity Poor Camarasaurus

Articulated skeleton of juvenile Camarasaurus lentus in the Carnegie Museum of Natural History, photographed by Daderot.


Let me just get the obvious out of the way first: sauropods were huge. Mind-bendingly huge. In some cases, big enough to reduce a human to a sticky puddle under foot and not even break their stride. For close to 150 million years, they were the largest land animals anywhere in the world, and no other terrestrial animal at any time has come even close to rivalling their largest representatives in size. Being around for so long, it should also be no surprise that they were diverse: a large number of sauropod genera have been named, representing a wide variety of forms. Nevertheless, most people's idea of sauropods is encompassed within just four genera from the late Jurassic of North America: Diplodocus, Apatosaurus, Brachiosaurus and Camarasaurus.

These four genera all belong to the clade Neosauropoda, which has been defined as the smallest clade containing the genera Diplodocus and Saltasaurus (a late Cretaceous South American genus). Upchurch et al. (2004) diagnosed the neosauropods by a number of cranial features, together with a reduction in the fourth hind toe (part of a general trend towards toe reduction in sauropods as their feet became more columnar—see this article by Darren Naish for more on the subject). However, Upchurch et al. were writing before the recognition of the Turiasauria, a European clade that is probably the sister group of neosauropods (Royo-Torres et al. 2006), and I don't know how that clade would affect the synapomorphy distribution*. The neosauropods quickly became the dominant sauropod group after their appearance in the middle Jurassic, and the only non-neosauropod sauropods to make it into the early Cretaceous were the aforementioned turiasaurs and possibly Jobaria, an African genus that varying analyses place either just inside or just outside the Neosauropoda.

*There has been an annoying tendency in recent years for many papers featuring phylogenetic analyses to present us with the character data matrix and the final trees from the analysis, but not do anything to map character changes onto the tree. The only way to find that out would be by transcribing the entire matrix and re-running the analysis yourself.

Mounted skeleton of Amargasaurus cazaui in the Melbourne Museum.


The famed North American genera include representatives of each of the two main lineages within the Neosauropoda. Diplodocus and Apatosaurus belong the Diplodocoidea, and Brachiosaurus and Camarasaurus belong to the Macronaria. Diagnostic features of the diplodocoids according the Upchurch et al. (2004) include restriction of teeth to the front of the jaw and a subrectangular snout. This last feature reaches an extreme in the middle Cretaceous Nigersaurus, which was another of those animals that serves to remind us that, if God did indeed create all of nature, then he was taking the piss. Ludicrous diplodocoids also include the late Jurassic South American Brachytrachelopan, which took one look at the graceful, elongate necks of all the other sauropods and decided that it simply couldn't be having with all that.

Reconstruction of Saltasaurus loricatus, by Lady of Hats.


The name of the other lineage, Macronaria, means 'big nostrils', and one of the notable features of this clade is, indeed, a great expansion in the size of the nares, the opening for the nostrils in the skull (though whether the size of the nares directly corresponds to the size of the actual nostrils is, I suppose, another question). As well as Brachiosaurus and Camarasaurus, this clade includes the Titanosauria, a very successful group that included the last surviving sauropods, but whose significance was overlooked for many years because they had the poor judgement not to achieve their main diversity in North America. At least some titanosaurs, such as Saltasaurus pictured above, sported a skin reinforced by nodules of bone.

Which brings us to Camarasaurus. For some reason, of the 'Big Four' genera, this is the one that gets the least love. While the other three have each in their turn enjoyed roles as stars of stage and screen, I'm not aware of a single film in which Camarasaurus has even been given a name-drop*. In John Sibbick's illustration of Brachiosaurus and Camarasaurus together (scroll down a bit at the link) in Norman's (1985) The Illustrated Encyclopedia of Dinosaurs, Brachiosaurus marches confidently towards the front bearing a goofy leer, while Camarasaurus is forced to sulk towards the back. It's all blatantly unfair. It's not as if Camarasaurus is rare: in fact, Camarasaurus may just be the best known of all sauropods, represented in the Morrison Formation by a whole whack of remains, including what is perhaps the single most beautiful sauropod specimen ever found (the one with which I opened this post). It was no slouch in the size department, either: its maximum length of about 23 metres is similar to that of Apatosaurus, despite it having proportionally shorter appendages than the latter. Nor does it lack distinctiveness: the short, bulldog-like face of Camarasaurus instantly stands out in any neosauropod line-up. So after all this time, doesn't Camarasaurus deserve to be given the spotlight?

*Though it is a pity that the name 'Camarasaurus' won out in the priority stakes over its competitor 'Morosaurus', which to those of us from a New Zealand background suggests a dinosaur made out of chocolate.

REFERENCES

Norman, D. 1985. The Illustrated Encyclopedia of Dinosaurs. Salamander Books: London.

Royo-Torres, R., A. Cobos & L. Alcalá. 2006. A giant European dinosaur and a new sauropod clade. Science 314: 1925-1927.

Upchurch, P., P. M. Barrett & P. Dodson. 2004. Sauropoda. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria, 2nd ed., pp. 259-322. University of California Press: Berkeley.

Sleepers

Hawaiian sleeper Eleotris sandwicensis, from the Hawaii Biological Survey.


Fishes of the genus Eleotris are a group of gobioids commonly known as the spinycheek sleepers. I haven't found a definite statement as to why they're called sleepers, but presumably it's because, as sit-and-wait ambush predators, they spend a lot of time lying around on the bottom. Eleotris species are found in tropical and subtropical waters around the world, mostly in estuaries and freshwater. They are smallish fish, with most species seeming to be in the ten to twenty centimetre size range. The 'spinycheek' part of the vernacular name refers to the presence of a hook-like spine on the lower corner of the preoperculum (the bone running between the cheek and the gill cover on the side of the head). This spine may be covered with tissue and so not always readily visible, but Pusey et al. (2004) note that it 'can be easily detected by running a thumbnail lightly, and carefully, along the preoperculum margin'. Carefully, I think, is the operative word here.

Eleotris oxycephala, from Chinese Academy of Fishery Sciences.


The species of Eleotris are mostly a conservative bunch appearance-wise, and the genus seems to have gotten a reputation for being difficult to work with taxonomically (it doesn't help matters that for a long time 'Eleotris' was something of a dumping ground for generalised gobioids). The Japanese species were revised in 1967 by Akihito (yes, that Akihito), the West African species have been revised by Miller (1998), and the North and South American species by Pezold & Cage (2002), but species from the remainder of the Indo-Pacific remain unrevised. There has been some disagreement over the status of a group of New World species classified in the genus Erotelis, which resemble Eleotris species but are generally more elongate and have higher numbers of fin rays (Pezold & Cage 2002). Miller (1998) felt that this genus should be synonymised with Eleotris, but Pezold & Cage (2002) argued that its members were distinct enough to be kept separate. A molecular phylogenetic analysis of the gobioids by Thacker & Hardman (2005) suggested that 'Erotelis' is nested within Eleotris, which may support their synonymisation.

Dusky sleeper Eleotris fusca, photographed by C. Appleby.


The sleepers are amphidromous, meaning they spend part of their life in the sea. Sleepers enter the sea as larvae, returning to fresher waters as they mature. As a result of this marine stage in the life cycle, individual species of Eleotris may be widespread and can often be found in places such as oceanic islands that lack populations of permanently freshwater species. It has even been suggested they may cross oceans: Miller (1998), noting similarities between species on either side of the Atlantic, suggested that this may be the result of trans-Atlantic dispersal. Among the evidence cited in favour of this possibility was the record in 1987 of a specimen of the northern South American species Eleotris pisonis from the island of St Helena in the mid-Atlantic. However, Miller also noted that the amount of time it would take to disperse across the Atlantic is greater that the time it would take for the larva to develop to maturity (and mature Eleotris are not known from the open sea). Pezold and Cage (2002) were more skeptical about the possibility of trans-Atlantic dispersal, even though they admitted to being unable to identify any characters distinguishing the Caribbean E. amblyopsis from the West African E. daganensis. They queried whether the St Helena record may have been an individual transported in ship ballast water, rather than an unaided dispersal.

REFERENCES

Miller, P. J. 1998. The West African species of Eleotris and their systematic affinities (Teleostei: Gobioidei). Journal of Natural History 32 (2): 273-296.

Pezold, F., & B. Cage. 2002. A review of the spinycheek sleepers, genus Eleotris (Teleostei: Eleotridae), of the western hemisphere, with comparison to the West African species. Tulane Studies in Zoology and Botany 31: 19–63.

Pusey, B., M. Kennard & A. Arthington. 2004. Freshwater Fishes of North-eastern Australia. CSIRO Publishing: Collingwood.

Thacker, C. E., & M. A. Hardman. 2005. Molecular phylogeny of basal gobioid fishes: Rhyacichthyidae, Odontobutidae, Xenisthmidae, Eleotridae (Teleostei: Perciformes: Gobioidei). Molecular Phylogenetics and Evolution 37: 858-871.