Paramesochra acutata

Copepod taxonomy, it seems, is largely about counting setae. In his review of relationships within the interstitial harpacticoid family Paramesochridae, Huys (1987) recognised four species groups within the genus Paramesochra (which previously got a look-in at this site here). One of these groups, labelled the P. acutata-group, was characterised by reductions in numbers of setae, having lost the inner setae on the first segments of the endopods on the third and fourth legs.

Paramesochra taeana, a close relative of P. acutata, from Back & Lee (2010).

The group takes its name from the species Paramesochra acutata, described by Klie in 1935 from samples taken from coastal groundwater near the town of Schilksee on the northeastern coast of Germany, in the state of Schleswig-Holstein. Other notable features of P. acutata include the presence of four setae on the antennary exopod, well-developed narrow, triangular endopodal lobes on the modified fifth legs of the females, and conical caudal rami produced into spinose processes (Back & Lee 2013). I haven't been able to find whether P. acutata has been collected much beyond its initial locality but other members of its species group have been found around the world. One of these, P. hawaiensis, from (nach) Hawaii, is similar enough that it was until recently treated as a subspecies of P. acutata.

Appendages of female Paramesochra acutata, from Klie (1935).

So what, if anything, does all this mean? That, I'm afraid, is getting a bit beyond me. The fifth legs are used in spermatophore transfer and differences between species might presumably function in recognising suitable mates. Regarding the details of setation and ramus appearance, one wonders if there could be any relation to preferred micro-habitat. Are harpacticoids with fewer setae and more robust rami adapted for crawling among coarser sand grains? Honestly, I have no idea. Anyone care to find out?

REFERENCES

Back, J., & W. Lee. 2010. A new species of the genus Paramesochra (Copepoda: Harpacticoida) from Korean waters. Proceedings of the Biological Society of Washington 123 (1): 47–61.

Back, J., & W. Lee. 2013. Three new species of the genus Paramesochra T. Scott, 1892 (Copepoda: Harpacticoida: Paramesochridae) from Yellow Sea, Korea with a redescription of Paramesochra similis Kunz, 1936. Journal of Natural History 47 (5–12): 769–803.

Huys, R. 1987. Paramesochra T. Scott, 1892 (Copepoda, Harpacticoida): a revised key, including a new species from the SW Dutch coast and some remarks on the phylogeny of the Paramesochridae. Hydrobiologia 144: 193–210.

Klie, W. 1935. Die Harpacticoiden des Küstengrundwassers bei Schilksee (Kieler Förde). Schriften des Naturwissenschaftlichen Vereins für Schleswig-Holstein 20 (2): 409–421.

The Diosaccinae: Worldwide Sediment Dwellers

The harpacticoid copepods have been featured on this site a reasonable number of times now. These tiny crustaceans are among the most numerous animals in the world, both in terms of numbers of individuals and (in certain habitats) numbers of species. And among the most widespread representatives of the harpacticoids are members of the subfamily Diosaccinae.

Diosaccus tenuicornis, from Sars (1906).

The Diosaccinae are currently recognised as members of the family Miraciidae; earlier sources will usually refer to a family Diosaccidae but the recognition of the pelagic Miraciinae as derived members of this group (Willen 2000) requires use of the older name. Distinctive features of the Miraciidae compared to other harpacticoids include the presence of a relatively large, mobile rostrum and a number of distinctive arrangements of setae, including the inner seta on the basal endopodal segment of the first peraeopod (trunk leg) arising distally (Nicholls 1941, Willen 2000). Miraciids are also unusual in that females carry paired egg-sacs laterally; most other harpacticoid families carry only a single median egg-sac. Miraciids are divided between three subfamilies of which the Diosaccinae are the most diverse. Diosaccines are most readily distinguished by their retention of a number of plesiomorphic features such as crawling legs and relatively short caudal rami (Nicholls 1941; this author divided the current diosaccines between two subfamilies, the Diosaccinae sensu stricto and Amphiascinae, based on the presence or absence, respectively, of a clear distinction in breadth between the metasome and urosome, or 'trunk' and 'abdomen', but this division does not appear to have been recognised at this level by any subsequent authors). The great majority of diosaccines are marine, free-living and benthic. A handful of species have been described as associates of lobsters, whether commensals or semi-parasites. A small radiation of species of the genus Schizopera is known from Lake Tanganyika, and Karanovic & Reddy (2004) described a species Neomiscegenus indicus from subterranean fresh water in India. Marine diosaccines are found at all depths from the intertidal zone to the deep abyss. I don't know for sure but, though they are sediment dwellers, I don't get the impression (I could be wrong) that they are strictly meiofaunal. As noted earlier, many do not have the vermiform body shape characteristic of interstitial copepods. Many species also are around the half-millimetre size range, which I think may be relatively large for meiofauna?

Four species of Schizopera collected from Korea, from Karanovic & Cho (2016). Left to right: S. yeonghaensis, S. daejinensis, S. gangneungensis, S. sindoensis.

The other two subfamilies of Miraciidae are the aforementioned Miraciinae and the Stenheliinae, which have the endopod of the first peraeopod adapted for swimming rather than grasping and longer caudal rami. Though potential synapomorphies of the Diosaccinae were identified by Willen (2000), they're a bit weaksauce. There is a distinct possibility that further studies may identify the diosaccines as paraphyletic to the other two subfamilies. In particular, some diosaccines say a very unusual form of nauplius larva with the Stenheliinae, in which the body is strongly foreshortened and crab-like (Dahms et al. 2005). These nauplii also move sideways in a crab-like fashion and do not swim in the water column like the nauplii of other species. Practical considerations have lead most investigators of crustacean phylogeny to emphasis adult over larval morphology but the larval morphology of diosaccines raises some interesting questions.

REFERENCES

Karanovic, T., & Y. R. Reddy. 2004. A new genus and species of the family Diosaccidae (Copepoda: Harpacticoida) from the groundwaters of India. Journal of Crustacean Biology 24 (2): 246–260.

Nicholls, A. G. 1941. A revision of the families Diosaccidae Sars, 1906 and Laophontidae T. Scott, 1905 (Copepoda, Harpacticoida). Records of the South Australian Museum 7 (1): 65–110.

Willen, E. 2000. Phylogeny of the Thalestridimorpha Lang, 1944 (Crustacea, Copepoda). Cuvillier Verlag: Göttingen.

The Splanchnotrophidae: Comfy inside a Sea Slug

In previous posts, I've referred to the great significance of the minute crustaceans known as copepods to aquatic ecosystems. At the time, I was referring to free-living members of this group but the copepods also include a wide range of parasitic forms. Some of these parasitic copepods have evolved into forms so derived and bizarre that they are barely recognisable as crustaceans. One example of this is the family Splanchnotrophidae.

Sea slug Janolus fuscus with protruding egg sacs of a splanchnotrophid copepod, probably Ismaila belciki, copyright Michael D. Miller.

Splanchnotrophids are a group of copepods endoparasitic on two orders of shell-less marine gastropods (sea slugs), the Nudibranchia and Sacoglossa. They are characterised by reduced mouthparts and appendages though they retain a distinct pair of claw-like antennae. These antennae seem to be used to hold the copepod in place in their preferred location within the body cavity of their host. Though the exact means of feeding by splanchnotrophids is not certain, their rudimentary mouthparts, combined with a rarity of observations of actual tissue damage in parasitised hosts, indicate that they probably suck nutriment from their host's haemolymph. Females and males live in association within the host, the minute (and slightly more recognisably copepod-y) males holding close to their comparatively gigantic mates. As well as their size, female splanchnotrophids differ from males in the possession of elongate, tubular dorsal outgrowths of the thorax. These are most commonly presumed to function to provide more space for the female's enlarged ovaries, though some have suggested additional functions such as maintaining position within the host, respiration or absorbing nutrients (Anton & Schrödl 2013). The female's tubular egg-sacs extend through an opening in the host's body wall to release eggs into the water column. Usually, these egg-sacs will emerge close to some outgrowth of the host's own body, such as gills or papillae, and may be coiled if relatively long; these measures presumably help protect the egg-sacs from external damage. How the released larvae find and colonise new hosts remains unknown but it is possible the antennules (the smaller second pair of antennae possessed by most crustaceans) are used to locate hosts chemically, with their reduced condition in adults the result of a halt to development once their purpose has been fulfilled.

Female (left) and male Ismaila aliena dissected out from host, from Anton & Schrödl (2013).

Relatively few splanchnotrophids have been recognised to date, maybe about a dozen species divided between five genera. A few other species that had earlier been included in the family on little more grounds than that they were endoparasites of gastropods were excluded by Huys (2001)*. A sixth genus and species Chondrocarpus reticulosus is of uncertain relationships. If correctly associated with the splanchnotrophids, it is of interest in parasitising a different group of sea slugs (the pleurobranchids) and in its massive size (growing to twelve millimetres vs only a few millimetres for females of the other genera), but the only available description is inadequate for its proper characterisation. In some localities, splanchnotrophids have proven to be surprisingly abundant. A once-off survey of potential host species in Oregon found no less than 62% of individuals of one species to be infected (25 other potential host species were completely free of parasites), whereas a longer-term survey off the coast of Chile found an overall infection rate of 13% with some particular host species approaching 100% infection (Schrödl 2002). Host specificity seems to vary within the family: a study by Anton et al. (2018) found that species of the genus Ismaila tended to restrict themselves to a single host species, whereas species of Splanchnotrophus are more catholic and undiscriminating. Nevertheless, a lack of correlation between relationships of splanchnotrophid species and those of their host species suggests that, even in the more discriminating Ismaila, host changes may not have been uncommon.

*As a concise indication of just how sloppy some of the earlier work on 'splanchnotrophids' had been, one misattributed species was re-identified by Huys (2001) as having been based on the detached head of a pelagic amphipod.

The broader relationships of splanchnotrophids within copepods also remain poorly understood. A phylogenetic study by Anton & Schrödl (2013) suggested that Splanchnotrophidae may form a clade with another genus of copepods endoparasitic in gastropods, Briarella, with this clade being in turn derived from ectoparasitic ancestors. However, by the authors' own admission, this study was heavily biased in both taxon and character coverage to the Splanchnotrophidae, and may have been affected by insufficient scrutiny of non-splanchnotrophid taxa. Though derivation of the endoparasitic splanchnotrophids from ectoparasitic ancestors has a definite intuitive appeal, further study is required before we can feel confident about it.

REFERENCES

Anton, R. F., D. Schories, N. G. Wilson, M. Wolf, M. Abad & M. Schrödl. 2018. Host specificity versus plasticity: testing the morphology-based taxonomy of the endoparasitic copepod family Splanchnotrophidae with COI barcoding. Journal of the Marine Biological Association of the United Kingdom 98 (2): 231–243.

Anton, R. F., & M. Schrödl. 2013. The gastropod-crustacean connection: towards the phylogeny and evolution of the parasitic copepod family Splanchnotrophidae. Zoological Journal of the Linnean Society 167: 501–530.

Huys, R. 2001. Splanchnotropid systematics: a case of polyphyly and taxonomic myopia. Journal of Crustacean Biology 21 (1): 106–156.

Schrödl, M. 2002. Heavy infestation by endoparasitic copepod crustaceans (Poecilostomatoida: Splanchnotrophidae) in Chilean opisthobranch gastropods, with aspects of splanchnotrophid evolution. Organisms, Diversity & Evolution 2: 19–26.

Leptocaris: Living on the Edge

Some of the most remarkable faunal diversity in the marine environment is to be found in the interstitial spaces between grains of sand. Grazers, predators and scavengers can be found creating entire food webs at scales of less than one millimetre. The minute crustaceans known as copepods are among the more abundant inhabitants of the interstitial, and today's subject, Leptocaris, is among those interstitial copepods.

Dorsal habitus of female (left) and male Leptocaris ryukyuensis, from Song et al. (2012).

Leptocaris contains more than twenty-five species of extremely slender, cylindrical harpacticoid copepods growing to a bit over half a millimetre in length. Characteristic features of the genus include having the maxillipeds (one of the pairs of appendages making up the mouthparts) reduced or lost, and the proximal part of the endopod of the first swimming leg bearing a special anteriorly directed seta with a terminal comb (Song et al. 2012). Representatives of this genus have been collected from localities around the world though mostly in the Northern Hemisphere. Nevertheless, one can't help wondering how much of the genus' apparent rarity in the Southern Hemisphere is an artefact of low collection effort. This possibility should also be kept in mind when considering differences in the ranges of individual species: whereas many have only been collected from single localities (Song et al. 2012), the species L. trisetosus has been found from Finland to the Bahamas to South Africa, as well as in Korea with the last population being treated as a distinct subspecies (Lee & Chang 2008).

The majority of collections of Leptocaris have been from among sand but the genus has also been found in other microhabitats. In general, they are found in sediments with a high organic content. They are found in euryhaline and eurythermal habitats: that is, locations subject to wide variations in salinity and temperature. These may include beaches and brackish pools. They have been found among decomposing leaves in mangrove swamps (offhand, I haven't found if the diet of Leptocaris has been firmly established but I suspect they are probably detritivores). One species, L. kunzi, was described from an estuarine lake in Louisiana; another, L. stromatolicolus, is known from among stromatolites in Mexico. Two species, L. brevicornis and L. sibiricus, have even been found in continental fresh waters in Europe as well as in coastal brackish waters (Song et al. 2012). Overall, Leptocaris species seem to be most abundant in marginal habitats that may be too harsh and unstable for other copepods, making them fronteir harpacticoids.

REFERENCES

Lee, J. M., & C. Y. Chang. 2008. Copepods of the genus Leptocaris (Harpacticoida: Darcythompsoniidae) from salt marshes in South Korea. Korean Journal of Systematic Zoology 24 (1): 89–98.

Song, S. J., H.-U. Dahms & J. S. Khim. 2012. A review of Leptocaris including a description of L. ryukyuensis sp. nov. (Copepoda: Harpacticoida: Darcythompsoniidae). Journal of the Marine Biological Association of the United Kingdom 92 (5): 1073–1081.

Life in Sand

Paramesochra mielkei, from Huys (1987).

Paramesochra is a genus of minute marine copepods found around the world. Over twenty species are currently assigned to the genus, but it is likely that many more await description. The extremely small size of paramesochrids (most are less than half a millimetre in length) reflects the interstitial habitat of most species described to date, i. e. they live among the grains of sand beneath the surface of their substrate. Also related to their choice of habitat is their vermiform (worm-like) shape and reduced setation compared to other copepods. These features also mean that they would be poor swimmers so they probably do not often emerge above the substrate surface. Most of the species described so far are from shallower waters, but this possibly reflects a lack of study of deep-sea species rather than reflecting true diversity. For instance, a survey of deep-sea Paramesochridae in the southern Atlantic and Antarctic Oceans by Gheerardyn & Veit-Köhler (2009) identified four species of Paramesochra, none of which corresponded to previously described species. These species probably do not have the same lifestyles as the shallow-water interstitial species due to the deep-sea substrate being fine mud rather than sand. Vasconcelos et al. (2009) suggested that another deep-sea paramesochrid, Kliopsyllus minor, might burrow in fluid mud or live in the 'organic fluff layer' (wonderful words) on top of the sediment. Deep-sea Paramesochra would probably be similar.

For the most part, genera of copepods have generally been distinguished mechanistically—different genera have different combinations of key features (usually related to the number of setae or segments on appendages)—without an explicit consideration of how those characters relate to phylogeny. However, Huys (1987) did propose a phylogenetic arrangement for the genera of Paramesochridae in which he suggested that Paramesochra formed a clade with the genera Kliopsyllus and Kunzia on the basis of their possessing single-segmented exopodites on the antennae and mandibles. However, while his tree shows Paramesochra as a monophyletic sister group to a clade of the other two genera, he did not identify any synapomorphies for Paramesochra. Instead, the features distinguishing it from the other two genera (two-segmented endopodites on the second to fourth legs, four setae on the distal exopodite segment of the first leg and two setae on the distal exopodite segment of the fourth leg) are resolved as plesiomorphies relative to the other clade. So if any of you feel inspired to spend your time dissecting and examining the legs of animals about 0.3 of a millimetre in total length, I know a potential research project going begging...

REFERENCES

Gheerardyn, H., & G. Veit-Köhler. 2009. Diversity and large-scale biogeography of Paramesochridae (Copepoda, Harpacticoida) in South Atlantic Abyssal Plains and the deep Southern Ocean. Deep-Sea Research I 56: 1804-1815.

Huys, R. 1987. Paramesochra T. Scott, 1892 (Copepoda, Harpacticoida): a revised key, including a new species from the SW Dutch coast and some remarks on the phylogeny of the Paramesochridae. Hydrobiologia 144: 193-210.

Vasconcelos, D. M., G. Veit-Köhler, J. Drewes & P. J. Parreira dos Santos. 2009. First record of the genus Kliopsyllus Kunz, 1962 (Copepoda Harpacticoida, Paramesochridae) from Northeastern Brazil with description of the deep-sea species Kliopsyllus minor sp. nov. Zootaxa 2096: 327-337.

Another Case of Mistaken Identity

Just the other day, Adam Yates showed us a couple of photos of a fossil that had been identified as dinosaurian, but actually belonged to a fish. Identifying isolated pieces of things can be a hazardous activity, and a mistaken identification can become something of a self-fulfilling prophecy - once the idea of a certain identity for your specimen has developed, you will tend to find "characters" that support your identification. Palaeontology, of course, presents researchers with no shortage of fragmentary remains, and it is not entirely surprising that a few snafus have occured. Adam referred to the case of Aachenosaurus multidens, a "hadrosaur" described in 1888 that was soon reidentified as a piece of petrified wood. A similar fate befell the "sauropod jaw" Succinodon putzeri (making the first four letters of the species name even more apropos). But while the most famous (and most dramatic) examples of such misidentifications involve fossils, studies of recent organisms have not been entirely free of impostors.



The figure above from Huys (2001) shows two views of the paratype of Megallecto thirioti, described by Gotto in 1986. The two specimens originally assigned to this species came from a plankton haul off the coast of Mauretania. Gotto identified them as parasitic copepods belonging to the family Splanchnotrophidae, and suggested that their hosts might be pteropods from the same haul.

Parasitic copepods can certainly be very strange creatures. While free-living males (and larvae of both sexes) may look like fairly ordinary copepods, the parasitic females may have highly derived morphologies that barely resemble crustaceans, let alone copepods. Consider the female of another splanchnotrophid, Arthurius elysiae (also from Huys, 2001):



When Huys (2001) revised the Splanchnotrophidae, however, he discovered that Gotto's Megallecto was (A) not a splanchnotrophid, and (B) not even a copepod. In fact:



'Megallecto' was nothing but a large chunk of the detached head of Phrosina semilunata, a pelagic amphipod. Phrosina belongs to a group of amphipods known as Hyperiidea. Most hyperiids feed on gelatinous plankton such as jellyfish or salps. They may or may not feed on pteropods.

REFERENCES

Huys, R. 2001. Splanchnotrophid systematics: A case of polyphyly and taxonomic myopia. Journal of Crustacean Biology 21 (1): 106-156.

Taxon of the Week: Some Copepods for your Reading Pleasure

The copepods are a widespread group of aquatic crustaceans, another one of those groups of minute animals that are all around us, yet attract little attention because of their tiny size. Thousands of copepod species have been described from every imaginable habitat involving a certain degree of water - thousands more doubtless remain to be described.

In the tiny world of copepods, members of the family Aegisthidae are relative giants, attaining massive sizes of more than 1.5mm in length (in contrast, the type specimen of their sister taxon, Romete bulbiseta, measures a mere 360 microns). Aegisthidae are one of the more basal members of the Harpacticoida, one of the largest orders of copepods. Basal members of the order are fusiform, but many of the more derived infaunal species have a more vermiform shape. Some three thousand harpacticoid species have been named to date, and they are second only to nematodes in abundance in the meiofauna (Seifried, 2003).

The relatively large size of the Aegisthidae is probably connected to their different lifestyle from other harpacticoids - rather than being epibenthic (living on the surface of the substrate) or infaunal (burrowing within the substrate), many Aegisthidae are hyperbenthic - that is, they live in the water column just above the surface of the substrate (there is a bit of confusion about correct terminology - other authors refer to "demersal zooplankton" or "benthopelagic plankton". Funnily enough, the choice of terminology is generally connected to which marine setting, whether tropical, temperate or deep sea, the author is mostly working with - Mees & Jones, 1997). There is also a tendency to lengthen the body shape, particularly the caudal rami, two spine-like extensions from the posterior end of the abdomen (the image of Aegisthus at top left of this post, from here, shows just how incredibly long the rami can get). A few species of Aegisthidae have gone the whole pelagic hog and become genuine members of the upper zooplankton, some of the relatively few harpacticoids (members of only three families) to have done so.

I had intended to centre this post on just one of the aegisthid subfamilies, the Cerviniopsinae. It is worth noting that Aegisthidae once referred to a much smaller collection of species, the current subfamily Aegisthinae (including the holoplanktic species). However, Seifried & Schminke (2003) argued that the 'Aegisthidae' of previous authors represented a derived subgroup of the previous family Cerviniidae, rendering the latter paraphyletic and calling for its sinking. The 'cerviniid' subfamily Cerviniopsinae was recognised as probably also paraphyletic with regard to Aegisthinae, and in particular a connection was suggested to the 'cerviniopsine' genus Pontostratiotes. Nevertheless, Seifried & Schminke did not formally remove the Cerviniopsinae from their classification, retaining it as a provisional grouping pending a proper phylogenetic analysis of the Aegisthidae. Most of the Cerviniopsinae do not appear to have been dealt with since Lang's major monograph of the harpacticoids back in 1948. Members of the Aegithinae share at least one distinguishing feature of the Cerviniopsinae, having the caudal furci opposed to each other rather than divergent as in the Cerviniinae (Montagna, 1979). Members of the Aegisthinae show a tendency to reduction of the mandibles in the males, with the adult males of some species such as Andromastax muricatus and Aegisthus mucronatus being entirely non-feeding.

REFERENCES

Mees, J., & M. B. Jones. 1997. The hyperbenthos. Oceanogr. Mar. Biol. Ann. Rev. 35: 221-255.

Montagna, P. A. 1979. Cervinia langi n. sp. and Pseudocervinia magna (Copepoda, Harpacticoida) from the Beaufort Sea (Alaska, USA). Transactions of the American Microscopical Society 98 (1): 77-88.

Seifried, S. 2003. Phylogeny of Harpacticoida (Copepoda): Revision of “Maxillipedasphalea” and Exanechentera. Cuvillier Verlag: Göttingen.

Seifried, S., & H. K. Schminke. 2003. Phylogenetic relationships at the base of Oligoarthra (Copepoda, Harpacticoida) with a new species as the cornerstone. Organisms, Diversity and Evolution 3: 13-37.