Sending Forget-me-nots

The Chatham Islands forget-me-not Myosotidium hortensia, from here.

I haven't been able to prepare a full post lately as we're currently in the field conducting our next survey round for the day job. In the meantime, I'll just content myself with a brief introducion to the Cynoglosseae. This is a tribe in the plant family Boraginaceae, redefined by Långström & Chase (2002) on the basis of molecular phylogeny to effectively correspond to the clade of Boraginaceae with heterocolpate pollen, as well as an undivided style with a single stigma (another tribe of Boraginaceae, the Boragineae, was covered in an earlier post). In the heterocolpate pollen of Cynoglosseae, the three apertures found in the pollen of other Boraginaceae alternate with an equal number of 'pseudoapertures'. The pseudoapertures represent gaps in the outer exine coat of the pollen grain like the apertures, but lack certain other features of the latter such as a concentration of cytoplasmic vesicles, as well as being longer and narrower (Hargrove & Simpson 2003).

Flower of camelbush Trichodesma zeyanicum, photographed by Ethel Aardvark.

Perhaps the most familiar members of this usage of Cynoglosseae are the forget-me-nots of the genus Myosotis, with other members including the hound's-tongue Cynoglossum officinale and, here in Australia, the camelbush Trichodesma zeylanicum. Offhand, camelbushes are generally one of the more prominent flowering plants here on Barrow Island, my current location, though they're one a bit of a low right now. There has been a bit of rain, and camelbush doesn't like to get its feet wet.

The traditional associations of forget-me-nots, of course, are right there in their name. There are a number of stories supposedly explaining how these flowers came to be associated with the memory of loved ones (surely the most ridiculous being the one that apparently has a knight drowning under the weight of a bouquet of the things) but the true reasons are probably lost to history. My own suspicion is that it is perhaps ultimately because forget-me-nots are relatively unassuming as flowers go, making them an ideal symbol of beauty that should not be overlooked for the sake of more flashy but perhaps less reliable competitors.

Hound's-tongue Cynoglossum officinale, from here. Native to Europe, this plant has become established in many parts of North America.

Hound's-tongue, on the other hand, seems to get its name from the resemblance of its leaves to its namesake. This plant doesn't seem to have quite the same hold on human affection as the forget-me-not, and the reason for this may be indicated by some of its other vernacular names: 'monk's nit' or 'beggar's lice', in reference to its sticky seeds that adhere to clothing (perhaps 'gypsy flower' derives from the same source?) and, even more damning, 'rats and mice', referring to its unmistakeable smell.

REFERENCES

Hargrove, L., & M. G. Simpson. 2003. Ultrastructure of heterocolpate pollen in Cryptantha (Boraginaceae). International Journal of Plant Sciences 164 (1): 137-151.

Långström, E., & M. W. Chase. 2002. Tribes of Boraginoideae (Boraginaceae) and placement of Antiphytum, Echiochilon, Ogastemma and Sericostoma: a phylogenetic analysis based on atpB plastid DNA sequence data. Plant Systematics and Evolution 234: 137-153.

The Nostocaceae: Tangled Filaments

Macroscopic growth of the cyanobacterium Nostoc commune, from here.

The cyanobacteria, commonly referred to as the 'blue-green algae', were one of the first groups of bacteria to be recognised as distinct. As our knowledge of bacteria has improved over the years, the distinctiveness of cyanobacteria continues to be supported: they are the only bacteria to contain chlorophyll, capturing energy from the light through the oxidation of water. Unfortunately, this confidence in their separation has not necessarily been carried over at lower levels. Many cyanobacterial 'families' and 'genera' have not been supported by more recent molecular analyses. Nevertheless, one clade that has been well supported is the nitrogen-fixing cyanobacteria, Hormogoneae.

Morphology-based classifications of cyanobacteria started, firstly, with the question of whether a species existed as independent cells, or whether they formed thread-like chains. The Hormogoneae are all chain-forming species, with the chain contained within a but they are also distinguished by the formation of heterocysts, large morphologically distinct cells within the chain that specialise in nitrogen fixation. Heterocysts are so specialised that they are unable to photosynthesise for themselves and are dependent on their neighbour cells for nutrition. So, with the presence of differentiated, interdependent cells, Hormogoneae can be regarded not simply as colonies of cells but as true, multicellular bacteria. A number of species of Hormogoneae (particularly in the genus Nostoc) form symbiotic associations with plants that take advantage of their nitrogen-fixing properties. One of the best-known examples is the association of the aquatic floating fern Azolla with the cyanobacterium 'Anabaena' azollae, and Azolla is used as a source of nitrogen in rice paddies. In another post, I have described the association between a Nostoc species and the plant genus Gunnera.

Trichomes of Anabaena, from here. The yellowish cells are heterocysts.

Within the Hormogoneae, classifications have traditionally distinguished between the orders Nostocales and Stigonematales. Stigonematales have branching trichomes, while those of Nostocales are unbranched. The Nostocales have been divided between the Nostocaceae, Rivulariaceae and Scytonemataceae: Rivulariaceae have trichomes that show a distinct base-to-apex polarity, while those of Scytonemataceae show a feature called 'false branching'*. Nostocaceae are defined by the lack of these features: however, it should not be surprising that molecular studies have not supported a group defined solely by the absence of characters, and both Rivulariaceae and Scytonemataceae (and possibly Stigonematales as well) are probably derived (possibly polyphyletically) from 'nostocacean' ancestors. However, actual relationships within the Hormogoneae remain poorly resolved, and no formal reclassification has been proposed (one of the authoritative texts on bacterial classification, Bergey's Manual of Systematic Bacteriology, replaces the cyanobacterial 'orders' with numbered subsections [Nostocales, for instance, is treated as Cyanobacteria subsection IV]—Castenholz 2001).

*True branching as in Stigonematales occurs when cells within a trichome divide at right angles to the direction of the trichome. In 'false branching', the trichome breaks within the containing sheath and then grows out of the sheath, but the division of the individual cells remains linear.

Trichomes of Cylindrospermum licheniforme, from André Advocat. The heterocysts are the round terminal cells, while the large elongate cells behind them are akinetes.

Phylogenetic analyses have also failed to confirm many of the genera recognised within the Nostocaceae (treated by Bergey's Manual as 'form-genera' only), distinguished by features such as whether trichomes are generally straight or coiled, and the positions within the trichome of heterocysts and other specialised cells called akinetes, thick-walled cells that function as resistent spores. The traditional genus Nostoc also differs from other Nostocaceae by the formation of hormogonia, motile trichomes with smaller cells and without differentiated heterocysts. Hormogonia form the dispersal stage of the Nostoc life cycle; it is as hormogonia, for instance, that symbiotic Nostoc are transmitted to new hosts. Mature Nostoc trichomes are embedded in a gelatinous matrix, and in some species this matrix may form a globular ball containing large numbers of radially arranged trichomes. Though usually microscopic, these globular clusters can get very large, sometimes more than twenty centimetres in diameter. Species attributed to other genera of Nostocaceae do not generally produce differentiated hormogonia (though some do, such as the aforementioned Anabaena azollae). Trichomes of these species may remain motile throughout the life cycle, or they may be permanently immotile (the latter state is characteristic of planktonic species).

REFERENCE

Castenholz, R. W. 2001. Phylum BX. Cyanobacteria. Oxygenic photosynthetic bacteria. In: Boone, D. R., & R. W. Castenholz (eds) Bergey's Manual of Systematic Bacteriology 2nd ed., vol. 1, pp. 473-599. Springer.

Beaver Fever

Eurasian beaver Castor fiber, from here.

Beavers are one of those animals that are familiar even to people who do not live in parts of the world where you can find beavers. The two living species of beaver are semi-aquatic rodents with one species each native to Eurasia (Castor fiber) and North America (C. canadensis) (though the North American beaver has been introduced to several parts of Europe). Differences between the two are slight: the Eurasian beaver is generally larger (up to 35 kg) and has a somewhat longer skull and a less rounded tail. Beavers are best known, of course, for their construction of elaborate subaquatic nests and dams*. Dams are generally about fifteen to seventy metres across, but have been recorded over 600 metres across (Rybczynski 2008). Beavers may also dig burrows connected to their dams, and construct canals over one hundred metres long (Rybczynski 2008).

*The original text here has been edited following Howard's comment below.

Phylogenetically, beavers are somewhat remote from other rodents, and represent the last survivors of a once more diverse lineage. First known from the late Eocene, the members of the beaver family Castoridae are divided in the most recent treatments between five subfamilies (Korth 2001, 2004). The plesiomorphic subfamilies Agnotocastorinae and Anchitheriomyinae are not well known, and the Agnotocastorinae in particular may be non-monophyletic (Rybczynski 2007). The remaining subfamilies fall into two distinct lineages: one containing the Palaeocastorinae (Oligocene-Miocene), the other the Castoridinae (Oligocene-Pleistocene) and Castorinae (Oligocene-present). Of these two lineages, only the latter are known to have been semiaquatic: the Palaeocastorinae are strictly terrestrial.

Preserved Daimonelix burrow in the American Museum of Natural History, with specimen of Palaeocastor fossor in the presumed nesting chamber, photographed by Inazakira.

The palaeocastorines, a strictly North American lineage, were specialised burrowers. Their incisors, which have rounded faces in modern beavers, became flattened and adapted for digging. Their burrows were distinctive helicoidal structures, described as trace fossils under the name of Daimonelix ('devil's spiral'), that could reach over 2.5 metres in depth and twenty centimetres in diameter. These burrows were constructed in 'towns' with multiple burrows in close proximity. Though each burrow was independent, without connections between adjacent burrows, such close positioning suggests that palaeocastorines may have had a well-developed social structure (Hugueney & Escuillié 1996). However, though the palaeocastorines were much more diverse at their apogee than the castoroidine-castorine lineage, they became extinct after a relatively short period.

Reconstructed skeleton of Castoroides ohioensis alongside that of (I presume) a modern beaver in Earlham College, from here.

The Castorinae and Castoroidinae may never have achieved the diversity at any one point in time of the palaeocastorines, they were more successful over the long haul: the more diverse of the two subfamilies, the Castoroidinae, only became extinct fairly recently. Castoroidines are commonly referred to as the 'giant beavers', and while not all castoroidines were giant (many, if not most, were smaller than modern beavers), the largest of them certainly were: the North American Castoroides reached an estimated size of about 100 kg, and would have been as large as a small bear. Whether the giant Castoroides produced similarly gigantic dams, however, is uncertain. Evidence of wood-chopping behaviour like that known for modern beavers (in the form of preserved wood bearing identifiable tooth marks, in association with beaver remains) is only well supported for one fossil species, the castoroidine Dipoides (suggested evidence for wood-chopping in Castoroides is more equivocal) (Rybczynski 2008). Phylogenetic bracketing between Dipoides and modern beavers would suggest that wood-chopping arose at the base of the castoroidine-castorine clade; alternatively, the absence of direct evidence of such behaviour may suggest convergence between these two species. Also, Dipoides was a less efficient wood-cutter than modern Castor, cutting with the rounded edges of its incisors while Castor uses the flattened ends, and if it used chopped wood to construct nests then they would have probably been correspondingly more simple (beavers also use chopped wood for food, eating the leaves and bark, so wood-chopping does not automatically indicate dam-building). There are other indications that fossil beavers may not have been as specialised aquatically as the modern species: the early castorine Steneofiber, for instance, did not possess the flattened tail of Castor (a flattened tail has been indicated for Castoroides but Castor and Castoroides probably developed such tails independently) (Hugueney & Escuillié 1996).

Reconstruction of Trogontherium cuvieri, from Fostowicz-Frelik (2008).

Perhaps the primary enigma among fossil beavers is the European Pleistocene Trogontherium. Although also referred to as a 'giant beaver', and often implied to be a European parallel to Castoroides, Trogontherium was a quite different animal. Fostowicz-Frelik (2008) argued that leg proportions and other features indicate that Trogontherium was a more terrestrial, cursorial animal than other beavers (in particular, its narrowed rather than flattened toe bones suggest that it lacked the webbed feet of modern beavers). The phylogenetic analysis of beavers by Rybczynski (2007) placed Trogontherium as closely related to Castoroides, but certain plesiomorphies in its tooth morphology lead Rybczynski to suggest that this position was probably an artifact of convergences due to large size, and that Trogontherium should perhaps be in a much more basal position.

REFERENCES

Fostowicz-Frelik, Ł. 2008. First record of Trogontherium cuvieri (Mammalia, Rodentia) from the middle Pleistocene of Poland and review of the species. Geodiversitas 30 (4): 765-778.

Hugueney, M., & F. Escuillié. 1996. Fossil evidence for the origin of behavioral strategies in early Miocene Castoridae, and their role in the evolution of the family. Paleobiology 22 (4): 507-513.

Korth, W. W. 2001. Comments on the systematics and classification of the beavers (Rodentia, Castoridae). Journal of Mammalian Evolution 8 (4): 279-296.

Korth, W. W. 2004. Beavers (Rodentia, Castoridae) from the Runningwater Formation (Early Miocene, early Hemingfordian) of western Nebraska. Annals of Carnegie Museum 73 (2): 1-11.

Rybczynski, N. 2007. Castorid phylogenetics: implications for the evolution of swimming and tree-exploitation in beavers. Journal of Mammalian Evolution 14: 1-35.

Rybczynski, N. 2008. Woodcutting behavior in beavers (Castoridae, Rodentia): estimating ecological performance in a modern and a fossil taxon. Paleobiology 34 (3): 389-402.

Name the Bug # 58

It's been a while since we last had one of these, but I thought I'd put this up in preparation for tomorrow's post:

What kind of animal was this, photographed in a Polish forest? Because this is one of the easier ones, tell me the exact species, please. Attribution, as always, to follow.

The Athyrididae: Spiralia and Lamellae

A specimen of the Devonian Athyris fultonensis photographed by Kentuckiana Mike. This specimen has part of the shell broken away to expose the calcified spiralium underneath.

The Athyrididae were a family of brachiopods that lived from the Silurian to the Permian, or until the end of the Triassic if the Diplospirellidae and Retzioidea are derived from the athyridids (Alvarez et al. 1998). Many athyridids possessed concentric lamellae on the outside of the shell; in the Upper Devonian to Permian Cleiothyridina, these lamellae were developed into a dense forest of flat spines. Specimens of the Devonian species Athyris vittata with preserved colour patterns indicate that the presence of radial stripes (Blodgett et al. 1988).

The Athyrididae are members of the order Athyridida, one of a number of brachiopod groups to possess a calcified spiral support (called, funnily enough, a spiralium) for the lophophore (see the link above for an explanation of the brachiopod lophophore). In the 1965 Treatise on Invertebrate Paleontology volume for brachiopods (Moore 1965), all the spiralia-possessing brachiopods were combined as the Spiriferida; however, other features of the shell are not consistent with a single origin for the spiralium, and they are now divided between the Spiriferida, Atrypida and Athyridida.

The Lower Carboniferous lamellate athyridid Cleiothyridina sublamellosa, photographed by Dwergenpaartje. The fine spines projecting from the lamellae in this genus have been mostly worn off in this specimen.

Many spiralium-bearing species appear to have lived on soft sediments, and it is possible that the spiralium was developed primarily as an adaptation for such habitats (Alvarez & Brunton 1990). For filter-feeders living in such habitats, the greatest challenge for feeding is not so much taking in food particles, but keeping the filter (in this case, the lophophore) from becoming clogged by indigestible particles such as sand. Reversing the direction of beat of the lophophore cilia moves such particles back towards the shell opening, where rapidly closing the valves will give the final impetus to 'spit out' the offending particles. In those species with lamellae, the slow-down of water currents as they hit the lamellae before the water enters between the valves may have also reduced the amount of particulate matter getting in. The lamellae may have also helped to support the shell opening above the sediment surface, though more significant in this regard would have been the pedicle, the fleshy stalk emerging from an opening in the back of the shell that anchored the athyridid in the sediment.

REFERENCES

Alvarez, F., & C. H. C. Brunton. 1990. The shell-structure, growth and functional morphology of some Lower Devonian athyrids from northwest Spain. Lethaia 23: 117-131.

Alvarez, F., Rong J.-Y. & A. J. Boucot. 1998. The classification of athyridid brachiopods. Journal of Paleontology 72 (5): 827-855.

Blodgett, R. B., A. J. Boucot & W. F. Koch II. 1988. New occurrences of color patterns in Devonian articulate brachiopods. Journal of Paleontology 62 (1): 46-51.

Moore, R. C. (ed.) 1965. Treatise on Invertebrate Paleontology pt H. Brachiopoda, vol. 2. The Geological Society of America, Inc., and the University of Kansas Press.

The Saga of Forsteropsalis fabulosa

Male of Forsteropsalis fabulosa*, photographed by Neil Fitzgerald. (Update: This male is actually F. bona, not F. fabulosa (see here. The two species are very similar; features of 'F. fabulosa' pointed out below for this photo still apply.)

Technically, I had a paper last week. I say 'technically' because, at only one page long (excluding bibliography), I don't that it counts much in the grand scheme of things. This commentary might be longer than the paper itself. Still, in its own way, this is a resolution for something that's been hanging over me for the last ten years.

The article, published in Zootaxa, is titled Clarification of the type status of Macropsalis fabulosa Phillipps & Grimmett 1932. Macropsalis fabulosa is the harvestman species that currently goes by the name of Forsteropsalis fabulosa. It is one of the largest of New Zealand's harvestmen, with a body length (excluding legs and chelicerae) a little shy of a centimetre (Phillipps & Grimmett 1932). As you can see in the photo at the top of this post, the male has chelicerae that are massively enlarged even by the standards of the group that it belongs to. The second inflated segment of the chelicera is about as large as the main body of the animal!

When I looked at all the (available) type specimens of New Zealand Enantiobuninae for my Master of Science thesis back in 2001, I discovered a problem with the type specimen of M. fabulosa. When a new species is described, the specimen(s) on which the description is based becomes the holotype (if a single specimen is used or designated) or the syntypes (if there is more than one specimen). However, sometimes the original type material of a species may become unavailable: it may be lost, destroyed, or subsequent researchers may not be able to identify what specimens the original author was using. In these cases, it may be necessary for an author to designate a neotype, a replacement type specimen. This is what had happened for M. fabulosa: Ray Forster had designated a neotype for it in 1944 because the original holotype had been destroyed. However, when I looked at Forster's neotype and compared it with the illustration of the holotype in Phillipps & Grimmett's original description, I realised that the two were not the same species! Here is the neotype, taken from the Te Papa online collection:

Compare the size and shape of the cheliceral fingers in that photo to the one at the top of this post. Also, while it is not clear in the photo, the neotype has a heavy covering of small spines over the top of the prosoma (cephalothorax), but Phillipps & Grimmett had specifically noted that, despite its size, M. fabulosa had a prosoma devoid of spines.

This raised a problem: should the name 'Macropsalis fabulosa' be applied to the species represented in the original description, or to the species represented by the neotype? My own preference was towards the former option:

(1) Despite his choice of neotype, Forster (1944) had still maintained the characters of the original holotype in his verbal description of M. fabulosa. It is quite possible that his choice of neotype was a mistake.

(2) The neotype belonged to a named species, currently known as Forsteropsalis inconstans. Because fabulosa is an older name than inconstans, associating fabulosa with the neotype would mean that F. inconstans would have to be called F. fabulosa, while F. fabulosa would be unnamed.

However, to keep the name fabulosa with the original species would require replacing the neotype, and that is something that only the ICZN can do. So last year, myself and my supervisor at the Western Australian Museum, Mark Harvey, drafted an application to the ICZN asking that the neotype of M. fabulosa be replaced with one of the specimens I had attributed to that species in last year's paper on Forsteropsalis (Taylor 2011). The application was duly submitted, but returned a few weeks later with explanations from the commissioners who had reviewed it that the ICZN would not be considering this case. Because they didn't need to.

The ICZN, in its current form, has rather stringent requirements for establishing a neotype. Basically, you cannot designate a neotype for a species solely because it doesn't have a holotype. There has to be an actual need for one, i.e. the species would not be properly identifiable without one. You also have to demonstrate that you took all the steps you could to make sure that the original holotype is honestly, truly lost (many specimens, for instance, may end up in a different museum from the one that the original author said they were in). Forster hadn't done that, and he had also violated the requirements by selecting a specimen that didn't match the diagnostic features of the species. So, according to the commissioners, there was no need to ask the ICZN to replace Forster's neotype because it wasn't valid in the first place.

But without the 'neotype' to confuse matters, there was no need for me to designate a new neotype either. Phillips & Grimmett's original description is quite adequate to identify M. fabulosa (that was how I had identified specimens of it myself). So, after that long process, I end up writing an article explaining why I have nothing to explain. One interesting potential side-effect is that (as one of the commissioners explicitly noted) this case is not unusual: so stringent are the current requirements for neotypification, without any protection for past practices, that probably a great many past neotype designations are technically invalid. However, they remain unchallenged because, in the vast majority of cases, there is no actual need for a neotype.

REFERENCES

Forster, R. R. 1944. The genus Megalopsalis Roewer in New Zealand with keys to the New Zealand genera of Opiliones. Records of the Dominion Museum 1 (1): 183–192.

Phillipps, W. J., & R. E. R. Grimmett. 1932. Some new Opiliones from New Zealand. Proceedings of the Zoological Society of London 1932: 731–740.

Taylor, C. K. 2011. Revision of the genus Megalopsalis (Arachnida: Opiliones: Phalangioidea) in Australia and New Zealand and implications for phalangioid classification. Zootaxa 2773: 1–65.

Life Among a Shrimp's Gills

Female of Schizobopyrina bombyliaster from Williams & Boyko (2004), with red box added on ventral view to indicate position of small male.

For today's random subject, I drew the marine isopod genus Schizobopyrina. Schizobopyrina is a genus in the family Bopyridae, and females of this genus were distinguished by Markham (1985) from those of the related genus Bopyrina by the presence of palp on the maxilliped (part of the mouthparts), by its more elongate oostegites (the lamellae forming the brood pouch in which eggs and larvae are incubated), and by the fusion of the pleomeres (posterior segments) on one side of the body. About ten or so species have been assigned to this genus from warmer waters around the world.

Mature bopyrids are parasites of shrimps and other crustaceans (Schizobopyrina has been found on hosts of the families Palaemonidae, Gnathophyllidae and Hippolytidae). Schizobopyrina and related genera are found in the branchial (gill) cavities of their host. Shrimp gills are developed from side-branches of the base of the legs, and are covered by an overhanging shelf of the carapace (if anyone is familiar with the process of preparing a crayfish or lobster, the gills are the 'dead man's fingers' that you have to remove before serving the crayfish). In a shrimp that is host to Schizobopyrina, the branchial cavity will become greatly protruding, as can be seen in this photo of a bumblebee shrimp Gnathophyllum americanum parasitised by Schizobopyrina bombyliaster (from Williams & Boyko 2004; scale bar equals 1.0 mm):

Bopyrids are released from the parent host as larvae that initially attach themselves to copepods. When they are approaching maturity, they leave the copepod and find an appropriate adult host. The first larva to attach itself to an appropriate shrimp will develop into a female, while any subsequent larva to attach itself will develop into a male (Cash & Bauer 1993). As can be seen in the figure at the top of this post, the female is considerably larger than the male. She is also noticeably asymmetrical in her body form, though a single species may include individuals bent to either the left or the right (Markham 1985). The female bopyrid attaches herself to her host before it reaches maturity: this puts her at risk of losing her place as the host moults, but studies of another branchial parasite bopyrid, Probopyrus pandalicola, indicate that as the host cuticle tears away during the process of moulting, the female is able to reattach herself to the new cuticle underneath and keep her place (Cash & Bauer 1993). The smaller male looks very different to the female, and is much more symmetrical. He attaches himself to the female, but whether or how he feeds is unknown. In Probopyrus pandalicola, the female moults, then produces eggs, after each moult of her host; the male has been observed crawling at this point into the brood pouch of the female, where he presumably fertilises her eggs.

Just as a further aside, the recent description of the species featured in the figures used in this post, Schizobopyrina bombyliaster Williams & Boyko 2004, was of further interest because the type specimen of this parasitic isopod was itself host to a hyperparasitic isopod, the cabiropid Cabirops bombyliophila. Which gives me an idea for a matryoshka design...

REFERENCES

Cash, C. E., & R. T. Bauer. 1993. Adaptations of the branchial parasite Probopyrus pandalicola (Isopoda: Bopyridae) for survival and reproduction related to ecdysis of the host, Palaemonetes pugio (Caridea: Palaemonidae). Journal of Crustacean Biology 13 (1): 111-124.

Markham, J. C. 1985. A review of the bopyrid isopods infesting caridean shrimps in the northwestern Atlantic Ocean, with special reference to those collected during the Hourglass cruises in the Gulf of Mexico. Memoirs of the Hourglass Cruises 7 (3): 1-156.

Williams, J. D., & C. B. Boyko. 2004. A new species of Schizobopyrina Markham, 1985 (Crustacea: Isopoda: Bopyridae: Bopyrinae) parasitic on a Gnathophyllum shrimp from Polynesia, with description of an associated hyperparasitic isopoda (Crustacea: Isopoda: Cabiropidae). Proceedings of the California Academy of Sciences 55 (24): 439-450.