Field of Science

Antipatharia: The Black Corals

The black coral Antipathes, copyright Jez Tryner.

One piece of trivia I've learnt while looking stuff up for this post: the genus name Antipathes, from which the whole group of the Antipatharia derives its name, was coined to refer to the supposed ability of black coral to cure illnesses and protect against evil. It almost goes without saying that I found no indications that this evaluation was warranted.

The black corals of the Antipatharia are a group of colonial, sessile cnidarians that are found in marine waters around the world. They are predominantly deep-water animals, found mostly below the level of light penetrance. Those individuals that are found in shallower waters still keep to secluded habitats out of the light. Some of the shallowest communities are found in New Zealand at depths of only 4 m in the fiords of the South Island, where a rich concentration of tannins in the top layer of the water prevents light from reaching even that far down (Wagner et al. 2012). Black corals have been harvested in many parts of the world for jewellery (and also for their supposed curative properties referred to above), but they are very slow-growing animals. At least one colony subjected to radiocarbon dating was estimated to be over 4000 years old (Roark et al. 2009).

Wire coral Cirrhipathes, copyright Frédéric Ducarme.

Colonies of antipatharians may be highly branched, or they may form an unbranched whip (the latter forms are sometimes referred to as wire corals or whip corals). They may be only a few centimetres tall, or they may reach a length of several metres in the case of some wire corals (Wagner et al. 2012). The core of the colony is a stalk composed of chitin that varies in colour from jet black in the main stem to golden yellow at branch tips. The stalk is lined with spines that may be simple cones, or may be covered with denticles, or may even be branched and antler-like. In life, the stalk is encased in living tissue, so black corals are not actually black. Unlike other skeletonised cnidarians in which the polyps are recessed within the skeleton, those of antipatharians are entirely external to it. As a result, black corals are rarely found in locations where there is a lot of moving sediment in the water, as they lack the ability to entirely retract the polyps to protect them from abrasion. The individual polyps are usually only a few milimetres wide and up to a few centimetres tall when extended. All antipatharian polyps have six tentacles and six primary mesenteries; depending on the species, there may also be four or six secondary mesenteries, though members of the family Cladopathidae lack secondary mesenteries altogether.

The most recent classification of the Antipatharia divides it between seven families, some of which have been recognised only very recently. Because their deep-water habitat makes the study of live colonies difficult, and many features of the minute polyps become obscured in preserved material, earlier classifications focused heavily on features such as the branching arrangement of the colony, or the morphology of the spines on the skeletal axis. However, these features may be influenced by environmental factors, and their significance may have been overestimated. For instance, a molecular phylogenetic analysis by Brugler et al. (2013) found that the unbranched wire coral genus Cirrhipathes was polyphyletic and not separated from the branched genus Antipathes. Nevertheless, Brugler et al. did find that the higher-level relationships within the Antipatharia were mostly concordant with morphology, including the distinction of the seven families. These relationships included a divergent position for Leiopathes, the only genus with six secondary mesenteries; a clade including the bathyal families Schizopathidae and Cladopathidae, in which the polyps are transversely elongated; a close relationship between the families Myriopathidae and Stylopathidae, with polyps that are not elongated and have relatively short, subequal tentacles; and an association of the families Antipathidae and Aphanipathidae, in which the sagittal tentacles tend to be quite elongate relative to the lateral tentacles. There was still, of course, room for investigation: one notable anomaly is that the type species of Antipathes, A. dichotoma, was identified as a member of 'Aphanipathidae' rather than 'Antipathidae'. If correct, this would mean that aphanipathids should be called antipathids, while antipathids would be... something else.


Brugler, M. R., D. M. Opresko & S. C. France. 2013. The evolutionary history of the order Antipatharia (Cnidaria: Anthozoa: Hexacorallia) as inferred from mitochondrial and nuclear DNA: implications for black coral taxonomy and systematics. Zoological Journal of the Linnean Society 169: 312-361.

Roark, E. B., T. P. Guilderson, R. B. Dunbar, S. J. Fallon & D. A. Mucciarone. 2009. Extreme longevity in proteinaceous deep-sea corals. Proceedings of the National Academy of Sciences of the USA 106 (13): 5204-5208.

Wagner, D., D. J. Luck & R. J. Toonen. 2012. The biology and ecology of black corals (Cnidaria: Anthozoa: Hexacorallia: Antipatharia). Advances in Marine Biology 63: 67-132.

A Spoonful of Lemba

Lemba or hill coconut Curculigo latifolia, from here.

The south-east Asian plant known as lemba has been referred to briefly on this site before, as a member of the family Hypoxidaceae. As noted in that post, it has been through a couple of names over the years: some sources will refer to it as Molineria latifolia, while others will call it Curculigo latifolia. The genera Molineria and Curculigo have been distinguished based on the presence of beaked (Curculigo) or unbeaked (Molineria) fruits and seeds, but the phylogenetic analysis of Hypoxidaceae by Kocyan et al. (2011) did not find this character to correlate with phylogeny. They therefore proposed to stop recognising the two genera as distinct, merging all species under Curculigo.

Curculigo latifolia is one of the largest species in the Hypoxidaceae. It is mostly found growing in damp, shaded locations, and the long-petioled leaves coming from an erect central rhizome can be a metre in length. Its small yellow flowers are placed at the base of the plant, at ground level; this distinguishes this species from various large orchid species found in the same region that may also be referred to as 'lemba' (or 'lumbah', or some other spelling/linguistic variant). The flowers give rise to small white berries, about an inch in size, with a distinct beak.

Fruit cluster of Curculigo latifolia, from DQ Farm.

Uses of this plant were recently reviewed by Lim (2012). The leaves provide a strong, lightweight fibre that is used to make nets, rope and cloth. The roots are brewed to treat various illnesses. However, the feature of this plant that has received the most attention in recent years is the fruit. These are edible, and are said to taste a bit like a sweet cucumber. The reason they have aroused interest, though, is that after eating one, anything else eaten within the next ten minutes or so will also taste sweet. This effect has been traced to a protein in the fruit, variously called neoculin or curculin, that has been reported to have several hundreds times the sweetness relative to weight of sucrose. Curculin has consequently been proposed as a potential low-calorie sweetener (to which I say, I'm sure it can't possibly taste worse than stevia), though one limitation is that the protein becomes denatured at temperatures above fifty degrees and loses its sweetening properties. As yet, though, it doesn't look like lemba sweetener has made it onto the commercial market.


Kocyan, A., D. A. Snijman, F. Forest, D. S. Devey, J. V. Freudenstein, J. Wiland-Szymańska, M. W. Chase & P. J. Rudall. 2011. Molecular phylogenetics of Hypoxidaceae—evidence from plastid DNA data and inferences on morphology and biogeography. Molecular Phylogenetics and Evolution 60 (1): 122-136.

Lim, T. K. 2012. Edible Medicinal and Non-Medicinal Plants, vol. 4. Fruits. Springer.

Book Review: The Amazing World of Flyingfish, by Steve N. G. Howell

In June of last year, I was standing on the deck of a ferry in Taiwan, headed for the island of Lüdao (commonly known as Green Island), keeping an eye out for any interesting sights. I was particularly intrigued by the seabirds that I kept seeing flying away from the ferry. For some time, I couldn't make out exactly what kind of bird they were: they were small, and flew very quickly. Most oddly, they never seemed to rise very far above the water; I kept waiting for one to get higher so that I could get a better idea of its shape, but every time I tried to keep an eye on one individual, it would seem to disappear, as if it had re-entered the water. Eventually, understanding dawned: what I was seeing were not birds at all, but flying fish.

Flyingfish (Exocoetidae) are prominent members of the pelagic ecosystem in tropical waters. For some tropical seabirds (actual birds this time, such as boobies or frigatebirds), they are among the primary source of food. Steve Howell, author of The Amazing World of Flyingfish (Princeton University Press, who were kind enough to send me a review copy), put together a guide to flyingfish after travelling from New Zealand to Australia on the Spirit of Enderby as part of a cruise that was primarily supposed to be for bird-watching. But, as Howell explains, "birds tend to be few in the blue equatorial waters (remember, it's a desert, even though it's full of water), and attention sooner or later shifts to flyingfish".

The Amazing World of Flyingfish is not a large book: all up, it barely makes it over 50 pages. But almost every one of those pages is adorned with spectacular photographs that capture the grace and variety of flyingfish. The images chosen work wonders in expressing the liveliness of their subjects. My favourite image technically doesn't even show the fish at all: on p. 16, a triptych of photographs showing the process of re-entering the water shows first the fish in flight, then closing its fins as it approaches the water's surface, and then simply the splash as it disappears below. The text, geared towards a younger or a lay audience, provides a general overview of flyingfish, with chapters given self-explanatory titles such as, "What is a flyingfish?", "How big are they?", "How do they fly?"

And yet, I also found Howell's book frustrating. The numerous different flyingfish varieties depicted are labelled with vernacular names largely of his own creation, such as Atlantic patchwing, sargassum midget, Pacific necromancer. Zoological names are, for the most part, not provided. As Howell explains, most field guides to marine fish are written for biologists or fisherman, and are oriented around identifying a specimen after it has been caught, often relying on features (such as scale counts) that are not discernible in photographs of live individuals. As a result, the identity of most of Howell's 'field varieties' remains uncertain. But then, in another section of the book, we are told that one juvenile morph "was examined genetically and proved to be a young Atlantic Necromancer" (capitalisation Howell's), implying that the zoological identity of this species, at least, is known.

As Howell points out, "there remains an unfilled niche for a field guide that portrays flyingfish as observers see them in the air". Howell has produced an attractive and engaging introduction to the world of flyingfish, and it should provide an inspiration to fill that niche.

Aequitriradites: The Mark of the Cretaceous

Diagram of Aequitriradites ornatus, from Upshaw (1963).

The Cretaceous period is best known in popular culture as the time of Tyrannosaurus and Triceratops, of Pteranodon and Quetzalcoatlus, of Elasmosaurus and mosasaurs. But it was also, in an arguably even more significant way, the time of Aequitriradites.

Aequitriradites is the fossil represented in the diagram at the top of this post. It is not very large: at its largest, the species shown above is about a tenth of a millimetre across (Upshaw 1963). It is, in fact, the spore from a liverwort. The vegetative parts of liverworts mostly do not have much of a fossil record, being soft and prone to decay, though long-time readers may recall a suggestion that this spotty fossil record was occasionally dramatic (alas, general opinion seems to have not been swayed). However, their spores are more resistent, and hence may be abundant as fossils. Because it is rarely possible to tell exactly which plant they came from, fossil spores (and pollen) are classified as form taxa, parallel to the classification of other plant fossils. Aequitriradites species are characterised by a membranous flange (a zona) running around the outside of the spore, together with a triradiate laesura pattern (the fissures that mark where the spore opens when it germinates) on one face. Depending on the species, the laesurae may be well-marked or faint. There may also be an opening in the spore wall at the apex of the face opposite the laesurae (Cookson & Dettmann 1961). It has been suggested that the otherwise unidentified liverworts that produced Aequitriradites spores were probably related to the modern liverwort order Sphaerocarpales, and Archangelsky & Archangelsky (2005) compared Aequitriradites to the spores of the aquatic genus Riella.

Alternate faces of specimens of Aequitriradites plicatus, from Archangelsky & Archangelsky (2005).

The abundance of plant spores and pollen is such that they are commonly used as 'index fossils', indicators of the age of the rock they are found in. Aequitriradites contains various species throughout the Cretaceous period (species assigned to this genus from the Triassic seem to have since been re-classified). Li (2014) identified the appearance of Aequitriradites spinulosus in the very latest Jurassic as one of the better indicators of the start of the Cretaceous period in the Qinghai-Xizang Plateau in China. Aequitriradites species seem to have been most abundant in the Early Cretaceous, becoming rarer in the Late Cretaceous. Establishing the latest appearance of a spore taxon in the fossil record can be difficult, because of the possibility of re-working (fossil spores being disassociated from their original deposit and re-buried in a later one), but non-reworked examples of Aequitriradites in the latter part of the Late Cretaceous were alluded to by Askin (1990). TLDR: If you've got Aequitriradites, you've got Cretaceous.


Archangelsky, S., & A. Archangelsky. 2005. Aequitriradites Delcourt & Sprumont y Couperisporites Pocock, esporas de hepáticas, en el Cretácico Temprano de Patagonia, Argentina. Rev. Mus. Argentino Cienc. Nat., n. s. 7 (2): 119-138.

Askin, R. A. 1990. Cryptogam spores from the Upper Campanian and Maastrichtian of Seymour Island, Antarctica. Micropaleontology 36 (2): 141-156.

Cookson, I. C., & M. E. Dettmann. 1961. Reappraisal of the Mesozoic microspore genus Aequitriradites. Palaeontology 4 (3): 425-427, pl. 52.

Li, J. 2014. Upper Jurassic and Lower Cretaceous palynological successions in the Qinghai-Xizang Plateau, China. In; Rocha, R., et al. (eds) STRATI 2013, pp. 1197-1202. Springer Geology.

Upshaw, C. F. 1963. Occurrence of Aequitriradites in the Upper Cretaceous of Wyoming. Micropaleontology 9 (4): 427-431.

Tortoise Sorting

Indian star tortoise Geochelone elegans, copyright P. G. Palmer.

As a whole group, the 'true' tortoises of the family Testudinidae are easily recognised, with a usually terrestrial habitus (though at least one species, the serrated hinge-back tortoise Kinixys erosa, is a capable swimmer), columnar legs with short heavy-clawed feet, and a relatively high carapace. But relationships within the tortoises have seen a bit of shuffling around in recent years, and nowhere has that shuffling been more obvious than in the genus Geochelone.

Many of you may know Geochelone as the genus including the giant tortoises of the Galapagos Islands and islands in the Indian Ocean, as well as species such as the radiated tortoise G. radiata of Madagascar and the geometric tortoise G. geometrica of South Africa. At its broadest, about half the world's tortoise species have been included in Geochelone. However, the genus has always been poorly defined, marked by its 'primitive' skull (Gerlach 2001), and the lack of features of other tortoise genera such as the plastral hinge of the Palaearctic Testudo species, or the rear-carapace hinge of African Kinixys species. For the most part, Geochelone was simply a home for the bigger tortoises.

Burmese star tortoise Geochelone platynota, copyright Kalyar Platt.

So it is hardly surprising that phylogenetic studies, whether morphological (Gerlach 2001) or molecular (Fritz & Bininda-Emonds 2007), have failed to support a broad Geochelone as a monophyletic group. As a result, recent authors have advocated recognising a number of separate genera: the Galapagos tortoises belong to the genus Chelonoidis, the Seychelles giant tortoises to Aldabrachelys (as confirmed by ICZN 2013), the radiated tortoise to Astrochelys and the geometric tortose to Psammobates. From being the largest recognised tortoise genus, Geochelone has been cut down to a mere two or three species. These are the Indian star tortoise Geochelone elegans, the Burmese star tortoise G. platynota, and, maybe, the African spurred tortoise G. sulcata.

The Indian and Burmese star tortoises are two very similar species that get their names from their colour pattern, with radiating star-like markings on the carapace. The individual scutes of the carapace bulge outwards, giving the animal an overall lumpy appearance. They are both medium-sized tortoises, growing to over 30 cm in length. The Indian star tortoise Geochelone elegans is widespread in dry habitats in India, Pakistan and Sri Lanka, though conservation concerns have been raised about the extent of harvesting of wild tortoises for food and the exotic pet trade. The Burmese star tortoise G. platynota, on the other hand, is critically endangered, having been almost wiped out from its original range in the central dry zone of Burma. As long ago as 1863, Edward Blyth (who, offhand, sported one heck of an impressive beard) was complaining that specimens were difficult to find due to the local people's fondness for eating them (Platt et al. 2011).

African spurred tortoise Geochelone sulcata, copyright Chris Mattison.

The African spurred tortoise Geochelone sulcata is found across the southern part of the Sahara Desert and the Sahel. It is a particularly large tortoise, growing to over 80 cm (Swingland & Klemens 1989); in fact, it is the largest tortoise that is not found on oceanic islands. Spurred tortoises dig burrows that can reach up to 3.5 m in length in which to avoid the full heat of the day. This species is placed by some authors in its own genus Centrochelys, cutting Geochelone down to just the two Asian star tortoises. Molecular studies place G. sulcata as the sister taxon to the Asian species (Fritz & Bininda-Emonds 2007), but they have not been associated in morphological studies. Though widespread, the African spurred tortoise is regarded as vulnerable due to the degradation of its habitat. Concern has also been raised, again, about the collection of wild individuals for the pet trade.


Fritz, U., & O. R. P. Bininda-Emonds. 2007. When genes meet nomenclature: tortoise phylogeny and the shifting generic concepts of Testudo and Geochelone. Zoology 110: 298-307.

Gerlach, J. 2001. Tortoise phylogeny and the ‘Geochelone’ problem. Phelsuma 9 (Supplement A): 1-24.

ICZN. 2013. Opinion 2316: Testudo gigantea Schweigger, 1812 (currently Geochelone (Aldabrachelys) gigantea; Reptilia, Testudines): usage of the specific name conserved by maintenance of a designated neotype, and suppression of Testudo dussumieri Gray, 1831 (currently Dipsochelys dussumieri). Bulletin of Zoological Nomenclature 70 (1): 61-65.

Platt, S. G., T. Swe, W. K. Ko, K. Platt, K. M. Myo, T. R. Rainwater & D. Emmett. 2011. Geochelone platynota (Blyth 1863)—Burmese star tortoise, kye leik. Chelonian Research Monographs 5: 057.1-057.9.

Swingland, I. R., & M. W. Klemens (eds). 1989. The conservation biology of tortoises. Occasional Papers of the IUCN Species Survival Commission 5.

The Sweetest of Lips

Oblique-banded sweetlips Plectorhinchus lineatus, copyright Richard Ling.

In an earlier post on this site, I referred to a fish of the family Lethrinidae being known by the name of "sweetlips". However, as is usually the way with fish vernacular names, there is more than one family of fishes to which this name can be applied. 'Sweetlips' is also the vernacular name for fishes in the Plectorhinchinae.

The Plectorhinchinae is most commonly treated as a subfamily within the family Haemulidae, the grunts (some sources will place 'Plectorhynchidae' as a separate family, and no, that wasn't a typo: read on). Plectorhinchines are distinguished from other subfamily of haemulids, the Haemulinae, by characters including a longer dorsal fin and the presence of at least four prominent lateral line pores under the chin (Johnson 1980). The name 'sweetlips' refers to the prominent lips of mature individuals of two of the genera of plectorhinchines, Plectorhinchus and Diagramma, which are often a distinct colour from the rest of the head. Members of the third genus, Parapristipoma, have the lips not quite so prominent, and are commonly referred to as 'grunts' like the remaining haemulids.

African striped grunts Parapristipoma octolineatum, copyright Juan Cuetos.

The plectorhinchines are found around tropical reefs in the Indo-West Pacific and East Atlantic, with a single species, the rubberlip grunt Plectorhinchus mediterraneus, being found in the in the Mediterranean and Black Seas. No plectorhinchines are found on either side of the Americas. They are nocturnal predators of benthic invertebrates, emerging at night from the secluded crevices and overhangs where they spend the day. Most are medium-sized fish, though the painted sweetlips Diagramma pictum can get up to 90 cm. They are popular with fishers; Smith (1962) referred to them as "among the best if not the best eating fishes of the reef-haunting species". Many species can go through significant changes in coloration as they mature: spotted juveniles may become unicoloured adults, or blotchy babies may mature into stripes. The differences are great enough that juveniles and adults have often been mistaken for separate species.

Juvenile oriental sweetlips Plectorhinchus vittatus, copyright Jan Messersmith. The adult form of this species resembles the oblique-banded sweetlips in the top photo on this post.

But failure to associate parents with their children is not the only way in which this group has been dogged by confusing taxonomy. The name of the type genus has been variously spelled Plectorhinchus or Plectorhynchus, with the family name varying accordingly (it seems that 'Plectorhinchus' is the correct spelling). A surprising number of sources (e.g. Tavera et al. 2012) seem to have it both ways, with the genus being called Plectorhinchus but the higher taxon being called Plectorhynchinae (R. van der Laan et al. confirm the correct family-name spelling). Meanwhile, Smith (1962) argued for the use of the name Gaterin in place of Plectorhinchus, and called the family Gaterinidae. And if you have any interest in the vagaries of taxonomy, settle in: this is going to be a whole thing.

The name 'Gaterin' dates from what is usually known as Forsskål's (1775) Descriptiones animalium, which Fricke (2008) argued should be attributed to Niebuhr (see, right from the first sentence it's confusing). Peter Simon Forsskål and Carsten Niebuhr were members of a Danish scientific expedition in 1761 to 1763 to the Red Sea (though Forsskål himself was Swedish, but that's another story). Forsskål was the expedition's naturalist, while Niebuhr was there as a geographer. The history of the expedition, and of the composition of Descriptiones animalium, has been summarised by Fricke (2008). The expedition was particularly ill-fated; of six original members, Niebuhr was the only one to make it back to Denmark alive. After returning to Denmark, Niebuhr started preparing Forsskål's notes for publication. However, he found this no easy task. Forsskål had not prepared a single manuscript, but made notes on various scraps of paper; in the end, Niebuhr suspected that many of these scraps had gone missing. As an engineer, Niebuhr knew little Latin and even less biology, so he obtained the services of an academic adviser. The identity of this adviser was not divulged in the final publication by Niebuhr himself, but he has since been identified as the Danish naturalist Johann Christian Fabricius. The relationship between Niebuhr and Fabricius was not entirely positive (Niebuhr later stated that his adviser on Descriptiones animalium had been a 'strange fellow'), and Fabricius does not seem to have spent any more time on the Forsskål notes than he absolutely had to. As a result, the final publication that emerged was partly Forsskål, partly Niebuhr, partly Fabricius, and all dog's breakfast.

The name 'Gaterin' is listed by Forsskål/Niebuhr/Fabricius as one of the sub-divisions of the genus Sciaena, and Smith's (1962) revival of the name was based on the assumption that Forsskål intended these subdivisions to represent what we would now call subgenera. As such, Gaterin published in 1775 would clearly be an earlier name than Plectorhinchus published in 1802. Smith further supported this interpretation by pointing out that two names listed by 'Forsskål' as subdivisions of Chaetodon, Acanthurus and Abudefduf, had since been widely accepted as names for separate fish genera. There were no grounds, he claimed, for taking Abudefduf as valid but refusing Gaterin.

As it happens, Forsskål probably never intended either Gaterin or Abudefduf to represent generic names of any kind. It seems that his notes had used local Arabic names to refer to taxa to which he had not yet supplied formal Latin names. When Fabricius compiled these notes, he simply used the Arabic names as formal names, probably because he just didn't care. When 'Forsskål' referred to 'Gaterin' in his introductory paragraph for Sciaena, he was probably referring to the individual species known in Arabia as gaterin rather than any formal group. 'Abudefduf' may have been similarly inadvertent, but long usage as a genus name means that it should probably be retained whatever its original status. No such argument can be marshalled in favour of 'Gaterin', whose usage in place of Plectorhinchus has been minimal.

And I can think of no better response to all that than the expression of this painted sweetlips Diagramma pictum. Copyright John Natoli.


Fricke, R. 2008. Authorship, availability and validity of fish names described by Peter (Pehr) Simon Forsskål and Johann Christian Fabricius in the ‘Descriptiones animalium’ by Carsten Niebuhr in 1775 (Pisces). Stuttgarter Beiträge zur Naturkunde A, Neue Serie 1: 1–76.

Johnson, G. D. 1980. The limits and relationships of the Lutjanidae and associated families. Bulletin of the Scripps Institution of Oceanography 24: 1–114.

Smith, J. L. B. 1962. Fishes of the family Gaterinidae of the western Indian Ocean and the Red Sea with a resume of all known Indo Pacific species. Ichthyological Bulletin 25: 469-502.

Tavera, J. J., A. Acero P., E. B. Balart & G. Bernardi. 2012. Molecular phylogeny of grunts (Teleostei, Haemulidae), with an emphasis on the ecology, evolution, and speciation history of New World species. BMC Evolutionary Biology 12: 57.

The Eater of Light

The Waitomo harvestman Forsteropsalis photophaga, from Taylor & Probert (2014).

A little less than a year ago, I was contacted by a student at the University of Auckland in New Zealand, asking me about some harvestmen that she'd been trying to identify me from the Waitomo cave system. This incited a certain degree of excitement on my part, because I was not entirely unfamiliar with Waitomo's harvestmen. I had first seen specimens from there while doing my MSc back in 2001 or 2002, and had realised then that they represented an undescribed species. However, for various reasons, I had not yet published a description of the species in question. So when Anna contacted me, I decided it was time to bump the Waitomo harvestmen up the to-do list, and I replied to her asking if she would be interested in collaborating on a paper on the Waitomo harvestmen. She agreed, and the resulting paper came out just last week: C. K. Taylor & A. Probert, "Two new species of harvestmen (Opiliones, Eupnoi, Neopilionidae) from Waitomo, New Zealand".

Image of Waitomo cave, from here.

The Waitomo caves may be the world's only tourist attraction centred around an infestation of flies. The caves are home to an abundant population of glow-worms, larvae of the fungus gnat Arachnocampa luminosa. These fly larvae live on the roof of the cave, held in place by a silken hammock, and produce spots of brilliant blue light. It is the spectacle of these lights that draw the tourists, but for the glow-worms they serve a different purpose: the lights attract insects flying in the cave. In flying towards the light, insects become entangled in sticky threads that each glow-worm suspends below its hammock, providing the glow-worm with food. You can see a video of the process here, taken from the BBC's Planet Earth series.

But the glow-worms are not without predators of their own. With their long slender legs, harvestmen are able to carefully tip-toe between the sticky threads and pluck out the glow-worms, as from a luminous buffet (they also eat them at the pupal and adult stages). The harvestmen of Waitomo were studied by Myer-Rochow & Liddle (1988), who identified two species. One, a 'short-legged harvestman' Hendea myersi cavernicola (which actually has decidedly long legs, natch), is endemic to the cave system and was identified by Meyer-Rochow and Liddle as a strict troglobite (i.e. it spends its entire life within the cave). It has a number of features commonly associated with cave-dwelling, such as pale coloration and lengthened legs. It does differ from most troglobites in that it is not blind: while its eyesight is dim, it does retain enough to find its glowing prey.

Meyer-Rochow and Liddle also identified a 'long-legged harvestman' in the Waitomo caves, which they referred to as 'Megalopsalis tumida'. This name refers to a species first described from Wellington, quite some distance to the south, that now goes by the name of Forsteropsalis fabulosa (and has made an earlier appearance at this site, where it was the subject of this post). As it turns out, I identified two species of Forsteropsalis in material from the caves, neither of which was F. fabulosa. I can't be certain which was the species being looked at by Meyer-Rochow and Liddle. For various reasons, I suspect that they may have been looking at examples of both, but, as I've never been able to locate any vouchers for their study, I can't really say (remember, kids, vouchers are important).

Forsteropsalis bona, from Taylor & Probert (2014).

One of these species is indeed very similar to Forsteropsalis fabulosa, and has accordingly been labelled Forsteropsalis bona. Indeed, the two species are similar enough that I can now see that the photograph I used in my earlier post to illustrate F. fabulosa in fact shows an individual of F. bona. The primary difference between the two is in their pedipalps: in F. fabulosa, the patella of the pedipalp has a distinct finger-like process that is much reduced in F. bona. Forsteropsalis bona is not a strict troglobite: specimens have been collected at Waitomo both inside and outside the cave entrance. Instead, it is what is called a troglophile: individuals of F. bona probably use the caves as a cool, damp place to hang out during the day, emerging to forage outside the cave at night. This is the same pattern of behaviour found in New Zealand's cave wetas.

The second species is the beauty pictured at the top of this post. Its species name, photophaga, means 'eater of light', referring of course to its probable predation on glow-worms. This is a stunning animal: the enormous chelicerae typical of New Zealand Neopilionidae are rendered even more eye-catching by the presence of rows of longer spines (offhand, we don't yet know what the females of either of the Waitomo species look like, but they probably resemble other Forsteropsalis females in lacking the long chelicerae of the males). Whether Forsteropsalis photophaga is a troglobite or a troglophile is a bit more uncertain. I'm not aware of it having ever been collected outside the caves, but it doesn't seem to have the obvious modifications for cave-dwelling of Hendea myersi cavernicola (though when t comes to assessing elongated limbs in what is already a long-legged harvestman... how are you going to tell?). At present, I'm guessing troglophile rather than troglobite, but future studies may easily prove me wrong.

Forsteropsalis photophaga is also an intriguing animal from a taxonomic viewpoint. In the past, the two New Zealand harvestman genera Pantopsalis and Forsteropsalis have been pretty easy to distinguish, but F. photophaga has some features that are more reminiscent of Pantopsalis than of Forsteropsalis. Recently, other things have been brought to my attention that suggest that, while Pantopsalis as we currently know it still seems fairly robust, Forsteropsalis is beginning to look decidedly fuzzy around the edges. The relationship between these two genera (if, indeed, they should still be recognised as two separate genera) has still not been resolutely ironed out.


Meyer-Rochow, V. B., & A. R. Liddle. 1988. Structure and function of the eyes of two species of opilionid from New Zealand glow-worm caves (Megalopsalis tumida: Palpatores, and Hendea myersi cavernicola: Laniatores). Proceedings of the Royal Society of London Series B (Biological Sciences) 233: 293–319.