Field of Science

Stygophalangium: Harvestman or Mite?

The original illustration of Stygophalangium karamani, from Oudemans (1933).

In 1933, the Dutch zoologist Anthonie Oudemans described what he believed to be a remarkable new species of harvestman. Based on two specimens collected from an underground spring in modern-day Macedonia and dubbed Stygophalangium karamani, Oudemans regarded this as a highly degenerate form as a result of its habitat: small, soft-bodied, and eyeless. It exhibited some significant differences to other harvestmen: in particular, the body lacked obvious signs of external segmentation. Also, its apparent aquatic collection point stood in direct contrast to the otherwise terrestrial habitats of other species. Nevertheless, Oudemans placed this unusual animal in a new family, the Stygophalangiidae, and suggested that its reduced morphology compared to other harvestmen might be compared to the position of Eriophyes (a plant-feeding, four-legged genus) among the mites. However, due to its anomalous character, subsequent authors have not paid much attention to little Stygophalangium. Mello-Leitão (1944) briefly suggested that it might represent a primitive form, placing it at the base of a branch of the phylogenetic tree leading to the Cyphophthalmi (mite-like harvestmen) and Palpatores (long-legged harvestmen). A number of online sources, such as Wikipedia, refer to Stygophalangium as being classified with the Eupnoi (a subgroup of the Palpatores), but this claim seems to be baseless. It seems to be derived from Joel Hallan's online list of harvestman species (which no longer appears to be available) but while Oudemans did compare Stygophalangium to the eupnoin Phalangium opilio (the common field harvestman) in his original description, he did not actually classify his new species with any particular subgroup of harvestmen. Eventually, Kury (2011) dismissed Stygophalangium from consideration in his summary of harvestman classification, stating that it 'is probably a member of the Acari'.

Unfortunately, as much as Stygophalangium might not be a convincing harvestman, it is also not a very convincing mite. One of the primary features that lead Oudemans to see Stygophalangium as a harvestman was its possession of three-segmented chelicerae. Most arachnids have chelicerae with only two segments (the basal segment and an opposing mobile claw or fang); three-segmented chelicerae are only found in two groups, the harvestmen and the mite group Parasitiformes. Of the four main groups (Opilioacarida, Holothyrida, ticks and Mesostigmata) within the Parasitiformes, none are similar to Stygophalangium. The ticks have distinctly modified (and kind of terrifying) blood-sucking mouthparts. The Holothyrida and Mesostigmata are both armoured to varying degrees, and mesostigs also bear a branched structure called the tritosternum underneath the mouthparts that is not described for Stygophalangium. The Opilioacarida are large, superficially harvestman-like mites that also have visible indications of external segmentation. And while there are a number of known lineages of aquatic mites, none of them really looks anything like Stygophalangium. It would be surprising if Oudemans, one of the leading mite researchers of his time, failed to recognise a mite when he had one in front of him! It is true that Oudemans' work underwent a precipitous decline in his last years as a result of problems with his mental health (Southcott 1961), but at the time of Stygophalangium's publication Oudemans remained alert and well.

Ventral view of Stygophalangium, with close-ups of chelicera, terminal pedipalp segments, and leg claw, from Oudemans (1933).

So if Stygophalangium was not a harvestman, and not a mite, then what was it? It is possible, of course, that it represented some taxon that has never been recorded since, but such an agnostic interpretation simply leaves the question of its affinities open. We can still at least try and compare it to other animals as best we can. One quite important point that I have avoided mentioning so far is that Oudemans' specimens were apparently not mature: Oudemans was unable to find indications of either a genital or anal opening. Though he described the body as unsegmented, it should be noted that his illustration is a reconstruction of what was apparently a not so smoothly mounted animal. Oudemans did note that a number of creases were visible on the bodies of his specimens, though he interpreted these as artefacts of slide-mountaing rather than segment boundaries because they did not appear to be placed evenly (with some creases even crossing over each other). Also, the supposed aquatic habitat may be a red herring. Subterranean samples are commonly collected by lowering sampling devices down a borehole, and it is not unknown for surface-dwelling organisms to fall in the borehole or be picked up when the traps are raised or lowered. So is Stygophalangium a larval harvestman or mite?

Again, we can rule out any arachnid except harvestmen or parasitiform mites due to the three-segmented chelicerae. The objections given above to adult ticks or Mesostigmata apply equally well to their juveniles, so they're also out. Larval Holothyrida lack the heavy armour of the adults, but these large litter-dwelling mites are not found anywhere near Europe. On the harvestman side of things, most harvestmen as both adults and nymphs have the second pair of legs particularly long and filamentous, functioning in a similar manner to the antennae of insects. The only harvestmen to lack this feature are the Cyphophthalmi, and together with the Opilioacarida they are the only real candidates for comparison with Stygophalangium. Both are soil-dwelling animals, and both are known from the Balkan region.

Larva of Opilioacarus texanus, from Klompen (2000).

One point in favour of an opilioacarid identity is that Oudemans described the chelicerae of Stygophalangium as inserted more dorsally than in other harvestmen. Opilioacarids have similarly inserted chelicerae, with a hypostome extending underneath the chelicerae. Oudemans also described Stygophalangium as lacking setae dorsally (instead having a somewhat scaly texture); opilioacarids have dorsal setae on the prosoma only. The opiliacarid prelarva (the earliest stage of its life cycle) has a scaly texture very similar to Stygophalangium (Klompen 2000), but mite larvae and prelarvae have only three pairs of legs. If Stygophalangium is an opilioacarid, it would have to be one of the later nymphal instars in which the fourth pair of legs has developed. Other features of opilioacarid juveniles conflict with Stygophalangium, such as the two pairs of large eyes on the opilioacarid prosoma. Also, Oudemans illustrated the venter of Stygophalangium with the coxae (the basalmost leg segment) integrated with the underside of the body, whereas opilioacarids (like other Parasitiformes) have the coxae free from the venter and attached by sockets. As Oudemans indicated the coxae of Stygophalangium with dotted lines only, it is possible that he inferred their position under the assumption of harvestman affinities. However, even if we assume this to be the case and that what Oudemans took to be the trochanters (the second leg segment) were actually the coxae, then Stygophalangium is left with one leg segment too few.

Larva of Siro rubens, from Juberthie (1964).

The only information on the juvenile stages of Cyphophthalmi is a brief description of the larva of Siro rubens by Juberthie (1964). Cyphophthalmi lack obvious eyes, and their legs do have the right number of segments for Stygophalangium. Juberthie described the cyphophthalmid larva as lacking a developed anus, which correlates with Oudeman's description of Stygophalangium (opilioacarid nymphs, in contrast, have a well-developed anal cone). He also recorded the presence of a pair of egg-teeth in the midline of the prosoma near the front of the body, in the same position where Oudemans described a distinctive pigmented spot on Stygophalangium. Points against a cyphophthalmid identification include the non-dorsal insertion of the chelicerae (though, again, one can't help wondering about the possibility of distortion through slide-mounting) and the presence of sparse but distinct dorsal setae. Especially difficult are the pairs of large setae marking the positions of the repugnatorial tubercles on either side of the prosoma. Unfortunately, Juberthie did not describe the venter of the cyphophthalmid larva, or comment on the degree of sclerotisation (mature cyphophthalmids are heavily sclerotised, whereas Stygophalangium is explicitly soft-bodied).

And that is about as far as we can go without looking at the original specimens. Personally, I suspect the issues with a cyphophthalmid identification are easier to overcome than those with an opilioacarid one (perhaps Oudemans did indeed mistake segment boundaries for mounting artefacts, and perhaps the dorsal setae had been lost post-mortem and Oudemans overlooked their sockets) but any such judgement requires the original description to be at least partially erroneous. Oudemans said that his type specimens were deposited in the Rijksmuseum van Natuurlijke Historie in Leiden; I wonder if they're still there?


Juberthie, C. 1964. Recherches sur la biologie des opilions. Annales de Spéléologie 19 (1): 5–244.

Klompen, J. S. H. 2000. Prelarva and larva of Opilioacarus (Neocarus) texanus (Chamberlin and Mulaik) (Acari: Opilioacarida) with notes on the patterns of setae and lyrifissures. Journal of Natural History 34 (10): 1977–1992.

Oudemans, A. C. 1933. Ein neuer Stygobiont, Stygophalangium karamani Oudms. Zoologischer Anzeiger 103: 193–198.

Southcott, R. V. 1961. Studies on the systematics and biology of the Erythraeoidea (Acarina), with a critical revision of the genera and subfamilies. Australian Journal of Zoology 9: 367–610.

Serpularia: A Rightly Forgotten Problematicum

I think it may be time to rock out something that hasn't been seen on this site for a while. Horns at the ready...

(Credit, again, to Neil from Microecos). And I'm afraid that may just be the most excitement that we get in this post. While some fossils are problematic because they're so strange that they can't be easily compared to living animals, others are problematic simply because they're rubbish.

In 1840, the palaeontologist Georg Graf zu Münster ('Graf' being a German title that generally gets translated as 'Count') published his Beiträge zur Petrefakten-Kunde, in which he described a number of fossils held in his collection. This book included a section on fossils from the Ordovician Orthoceratite Limestone of the Fichtel Mountains in Bavaria. Which, close to the end, included this little tidbit:
Unter mehreren Bruchstücken einiger mir noch unbekannten Versteinerungen kommen auch einige röhrenformige Korper vor, welche ich anfänglich für den von Murchison aus der 27sten Tafel abgebildeten Myrianites hielt, allein genaue Untersuchung zeigte, dass diese Korper formliche Schalen hatten und daher vielleicht zu den Serpuliten gehört hatten, daher ich sie vorläufig Serpularia genannt habe. Aus der Taf. IX. Fig. 14 und 15 sind zwei Arten von dergleichen Bruchstücken abgebildet; Fig. II. Serpularia crenata; glatt gebogene Röhre, aus dem Rücken crenulirt. Fig. 15. Serpularia bicrenata; glatte etwas zusammengedrückte ganz grade Röhrchen, die an beiden Seiten crenulirt sind.

Translated with the help of Google Translate, I think this means: "Among several fragments of fossils unknown to me occured a tube-like body, which I initially took for Myrianites as figured by Murchison in the 27th plate, until close examination showed that this body had distinct signs of segmentation and was therefore perhaps one of the Serpulidae. Therefore, I have provisionally called it Serpularia. On Plate IX Figs 14 and 15 are shown two types of the like fragments; Fig. 14, Serpularia crenata: smooth curved tube crenulated from the back. Fig. 15, Serpularia bicrenata: smooth, slightly compressed, quite straight tubes that are crenulated on both sides".

Münster's (1840) original figures of the two Serpularia.

As perfunctory as it was, that seems to be all there was to say on the matter. The good Graf's Serpularia has pretty much never been mentioned again*, beyond being cited to cause a name change in a later homonymous gastropod genus, and a brief listing in Howell's (1962) coverage of worm fossils for the Treatise on Invertebrate Paleontology that adds nothing to the original description.

*Though if it were to be mentioned again, it would probably have to be under a different name. The name 'Serpularia' had earlier been used by Fries in 1829 for a genus of slime moulds. At the time, slime moulds were treated as fungi, and hence fell under the purview of botanical rather than zoological names, but with the recognition that they are amoebozoans an increasing number of authors would move them into the field of the Zoological Code.

Münster believed that his fossils belonged to the Serpulidae, a family of annelid worms. Annelids, being mostly soft and squishy things that do not stand up well to decay, have a pretty deplorable fossil record, but serpulids are a bit of an exception. These are sessile worms that secrete a calcareous tube in which they live their lives (modern serpulids appeared on this site in this post). Unfortunately, while these tubes are eminently fossilisable, they are also a bit nondescript, and have little to mark them as uniquely serpulid.

Because of the dominance of annelids among modern worms, there has been a definite tendency in the past to assume that any given worm-like fossil represents an annelid. Howell's (1962) aforementioned list of annelids includes the Ediacaran Spriggina (identity still under debate, but probably not an annelid) and the Cambrian Pikaia (now generally regarded as an early chordate). Similarly, any worm-like tube has been assumed a serpulid. But even among annelids, serpulids are not the only tube-bearing worms. At least two other families, the Sabellidae and the Cirratulidae, include species producing calcareous tubes. There are also other groups of non-annelid worms that, though relatively uncommon or unprepossessing today, may have been more prominent in the past. After all, we are talking here about a period of hundreds of millions of years. We know that vertebrates have gone through a great deal of evolutionary change over that period; why should we assume that worms have not?

So while fossils have been assigned to the serpulids going back as far as the Cambrian (if not beyond), there is little reason to take those assignations at face value. When so-called Palaeozoic serpulids have been examined critically in recent years, they have so far proven to lack features that would definitely confirm their identification (Vinn & Mutvei 2009). Weedon (1994) found that Palaeozoic fossils that had been assigned not only to the Serpulidae, but to the modern genus Spirorbis, had a shell microstructure that suggested a relationship to bryozoans or brachiozoans rather than to annelids. Without a similar close analysis, we could not assume a priori that Münster's Serpularia were not serpulids, but odds would currently be against it.


Howell, B. F. 1962. Worms. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt W. Miscellanea: Conodonts, Conoidal Shells of Uncertain Affinities, Worms, Trace Fossils and Problematica pp. W144–W177. Geological Society of America and University of Kansas Press.

Münster, G. 1840. Beiträge zur Petrefacten-Kunde von Herm. v. Meyer und Georg Graf zu Münster vol. 3. In Commission der Buchner'schen Buchhandlung: Bayreuth.

Vinn, O., & H. Mutvei. 2009. Calcareous tubeworms of the Phanerozoic. Estonian Journal of Earth Sciences 58 (4): 286–296.

Weedon, M. J. 1994. Tube microstructure of Recent and Jurassic serpulid polychaets and the question of the Palaeozoic 'spirorbids'. Acta Palaeontologica Polonica 39 (1): 1–15.

More on Spider-Hawks

A couple of years ago, I presented a bit of an abortive post on wasps of the family Pompilidae, the spider-hawks. Despite their striking appearance and relatively high visibility, I noted, it was nigh on impossible to find reliable taxonomic information on them.

Diagram of the forewings of Cryptocheilus australis (above) vs Heterodontonyx bicolor (below) from Wahis (2008), showing the differences in the shape of the marginal cell (the large cell along the top margin of the wing).

This question came back to the fore for me recently when I had to attempt to identify a number of spider-hawks for work. With no recent key available for Australian pompilids, I had to try and piece together clues. As it turns out, a large part of the difficulty in identifying spider-hawks is that they are, overall, a conservative bunch. Though coming in a range of sizes and colours, they tend to be structurally uniform. This makes it difficult to find reliably key-able characters, and means that evolutionarily quite distinct species can look superficially quite similar. Take, for example, one of the most 'familiar' of the Australian pompilids, the black-and-orange Cryptocheilus bicolor. Recently, Wahis (2008) established that this was not a true species of Cryptocheilus, but belonged to a distinct (albeit related) genus as Heterodontonyx bicolor. The two genera can be distinguished by the shape of the marginal cell in the forewing, which is distally pointed in Heterodontonyx but rounded in Cryptocheilus. The thing is, many of the photos one may find online labelled as 'Cryptocheilus bicolor' are true Cryptocheilus, not Heterodontonyx. Those on Wikipedia may be correctly identified, but these here are not. Not every large orange-and-black spider-hawk in Australia is Heterodontonyx bicolor.

Specimen of Telostegus inermis, copyright Josef Dvořák.

So what of Telostegus, the genus that I was complaining about being unable to find the diagnostic characters for in my earlier post? Evans (1972) describes it as having bifid tarsal claws, and a vena spuria in the forewing. A vena spuria ('spurious vein') is a fold in the wing that might be mistaken at first glance for a wing vein. In the images above, it can be seen as a dark line along the middle of the wing in the dorsal view. Evans (1972) separated two genera of spider-hawks, Telostegus and Elaphrosyron, on the basis of the number of submarginal cells in the forewing (two in Telostegus, three in Elaphrosyron) but more recent authors have not regarded this distinction as valid.


Evans, H. E. 1972. A review of the Australian species of Elaphrosyron and Telostegus, with notes on other genera (Hymenoptera: Pompilidae). Breviora 386: 1–18.

Wahis, R. 2008. Contribution à la connaissance des Pompilides d’Australie (Hymenoptera : Pompilidae). 2. Sur quelques spécimens récoltés par G. Else (Natural History Museum, London) avec descriptions de deux espèces nouvelles des genres Auplopus et Ctenostegus. Faunistic Entomology 61 (1–2): 23–31.

Hadromeros: A Trilobite Survivor

Reconstruction of Hadromeros subulatus, from Kielan-Jaworowska et al. (1991).

Trilobites of the genus Hadromeros were widespread in the Late Ordovician and Early Silurian of Eurasia and North America. They are classified in the Cheiruridae, the same family that includes another trilobite genus that has been featured on this site, Sphaerexochus. However, Hadromeros differs from Sphaerexochus in that its glabella (its 'nose') is not as large. Whereas in my earlier post I suggested that Sphaerexochus may have been a predator, Hadromeros was probably a less aggressive feeder. It was possibly a detritivore, picking bits of nutritious material out of the sand and mud. This interpretation is supported by the known leg morphology of a closely related genus, Ceraurus, in which the legs are fairly generalised and show little adaptation for food processing (Bergström 1973).

Hadromeros and Ceraurus are placed in the Cheirurinae, a distinct subfamily from the Sphaerexochinae that includes Sphaerexochus. One characteristic feature of many members of the Cheirurinae is that the bases of the pleura, the plates that run down each side of the thorax, are swollen in dorsal view. The purpose of these swellings remains unknown. You might expect that, trilobites being as abundant in the fossil record as they are, we would know a great deal about them, and in many respects that is quite true. However, in other respects our knowledge is also frustratingly incomplete. Trilobites have an extensive fossil record because their dorsal exoskeleton was mineralised, and it is this that is usually preserved. The ventral section of the body, on the other hand, was not mineralised, and is only preserved under exceptional circumstances. This includes such significant features as the legs and mouthparts (as indicated above, we have some knowledge of the leg morphology of Cerarurus, but no direct evidence for Hadromeros). It is possible that the cavities underneath the cheirurine pleural bases housed some modification of the gills, if the gills in trilobites were comparable to those of living crustaceans. But how or to what purpose the gills were modified can only be speculated upon.

Morphologically, Hadromeros was a fairly unremarkable trilobite, but it stands out from the other genera of the Cheirurinae in one important respect (that has, indeed, already been alluded to in this post). The end of the Ordovician saw a mass extinction in marine life, by some measures the second largest to have ever occurred. Few groups of animals made it through unscathed, and the cheirurids were no exception. Of the eight subfamilies of Cheiruridae recognised by Přibyl et al. (1985), only three made it through to the Silurian: the Cheirurinae, Sphaerexochinae and Deiphoninae. Within the Cheirurinae, Hadromeros is the only genus currently known from both sides of the Ordovician-Silurian boundary, and may have been ancestral to all other post-Ordovician cheirurines. While other genera were whisked away, Hadromeros became the Trilobite that Lived.


Bergström, J. 1983. Palaeoecologic aspects of an Ordovician Tretaspis fauna. Acta Geologica Polonica 23 (2): 179-206.

Kielan-Jaworowska, Z., J. Bergström & P. Ahlberg. 1991. Cheirurina (Trilobita) from the Upper Ordovician of Västergötland and other regions of Sweden. Geologiska Föreningen i Stockholm Förhandlingar 113 (2-3): 219-244.

Přibyl, A., J. Vaněk & I. Pek. 1985. Phylogeny and taxonomy of family Cheiruridae (Trilobita). Acta Universitatis Palackianae Olomucensis Facultas Rerum Naturalium Geographica-Geologica XXIV 83: 107-193.

The Dicranophyllales: An Early Branch of the Conifers?

Reconstruction of Dicranophyllum hallei, from here.

Popular works on the fossil record tend to give us a very uniform picture of the Carboniferous period. A watery swamp can be seen covering the landscape, from which large amphibians emerge onto sodden banks. Giant insects hover in the air. The vegetation is dominated by scaly-trunked lepidodendrons and enormous horsetails. The entire scene is primoeval, presenting us with the representatives of a generation of life long gone, whose like we shall never see again. But of course, not all of the Carboniferous world was given over to coal swamps. While the lepidodendrons and horsetails were indeed around, there were also the early representatives of more familiar plant lineages, though some of them may have been a bit difficult to recognise as such.

The Dicranophyllales may have been one such lineage. Though they survived for a long time, throughout the Carboniferous and Permian, and have been found in many parts of the world, they are generally uncommon in fossil deposits. In life, they would have been small trees or bushes, sparsely and irregularly branched (many reconstructions show them hardly branching at all). The branches bore long, needle-like leaves, not dissimilar to pine needles, in a helical arrangement. The longest of these leaves were over 20 cm in length. A single vein ran down the midline of the leaf, but because this was deeply imbedded it is often not visible in fossils. More prominent, and one of the characteristic features of the group, was a pair of deep grooves running the length of the leaf, one on each side close to the margin, containing the stomata (the openings through which planty leafs exchange gases with the surrounding atmosphere). The leaves were commonly branched towards the tips, at least once and sometimes more. The needle-like leaves, protected stomata, and uncommon preservation all suggest that the Dicranophyllales were mostly plants of drier environments (Wagner 2005). In many species, the leaves left a regular-shaped scar when they fell off, giving the trunk and branches an overall scaly appearance.

Reconstruction of a branch of Polyspermophyllum sergii, from Archangelsky & Cúneo (1990). Note the coiled fertile trusses at the ends of some leaves.

The majority of fossils of Dicranophyllales are of vegetative material (branches and leaves) only, and as a result they have mostly been assigned to the single genus Dicranophyllum, possessing the characters described above. Other genera of Dicranophyllales known from the Upper Permian of Russia include Mostotchkia, which differed in that the leaves were generally not branched, and Slivkovia, which had small scale-like leaves appressed to the branch surface in addition to the long needle-like leaves. Slivkovia and the Lower Permian Entsovia also differed from other Dicranophyllales in having a higher number of stomatiferous furrows on each leaf (Meyen & Smoller 1986). Reproductive structures are definitely recognised for only two species, the European Dicranophyllum gallicum, and Polyspermophyllum sergii from the early Permian of Argentina (Archangelsky & Cúneo 1990). Though Polyspermophyllum resembles Dicranophyllum vegetatively, it is distinct reproductively. In both species, the reproductive organs are broadly similar in appearance to the leaves, and occupy positions in the growth trajectory that would otherwise be occupied by leaves. Seeds are borne separately from each other on the female organs, which have been dubbed polysperms. In Dicranophyllum gallicum, the polysperms end in a bifurcation similar to that of a normal leaf, and the seeds are borne attached to the side. Unfortunately, the compressed fossils do not allow us to determine whether they were arranged helically or pinnately. The male organs were similar in organisation to the polysperms (Wagner 2005). In Polyspermophyllum, the polysperms are divided into multiple branches, and the seeds are borne in trusses at the ends of the branches.

Reconstruction of a section of Dicranophyllum gallicum bearing polysperms, from Seward (1919).

The affinities of the Dicranophyllales have been subject to debate. Some authors, such as Archangelsky & Cúneo (1990), have recognised two families in the Dicranophyllales: the Dicranophyllaceae containing all the taxa referred to above, and a second family including the Permian genus Trichopitys. Trichopitys is vegetatively similar to Dicranophyllales, but its leaves are arranged pinnately rather than helically, and its reproductive organs are borne axillary to the leaves rather than replacing the leaves in the growth sequence. As a result, other authors such as Meyen & Smoller (1986) have regarded the similarities between the two families as convergent. It has also been suggested that the Dicranophyllales might be early members of the lineage including the modern maidenhair tree Ginkgo biloba: under this model, the fan-shaped leaves of the ginkgo may be derived from branched leaves like those of Dicranophyllales by fusion of adjoining branches. However, Meyen & Smoller (1986) pointed out that the structure of Dicranophyllales leaves is less like those of a ginkgo that it is like those of early members of the conifer lineage. Some of the Cordaitanthales, a Palaeozoic group of plants related to the conifers, had furrows on their leaves similar to those found in Dicranophyllales. The leaves of Dicranophyllales also bear resemblances to those of early members of the conifers proper. And this is where the question of seed arrangement on the polysperms of Dicranophyllum becomes interesting: if they were helically arranged, then it becomes possible to the Dicranophyllum polysperm as a distant fore-runner of the modern pine cone.


Archangelsky, S., & R. Cúneo. 1990. Polyspermophyllum, a new Permian gymnosperm from Argentina, with considerations about the Dicranophyllales. Review of Palaeobotany and Palynology 63: 117-135.

Meyen, S. V., & H. G. Smoller. 1986. The genus Mostotchkia Chachlov (Upper Palaeozoic of Angaraland) and its bearing on the characteristics of the order Dicranophyllales (Pinopsida). Review of Palaeobotany and Palynology 47: 205-223.

Seward, A. C. 1919. Fossil Plants: A text-book for students of botany and geology vol. 4. Ginkgoales, Coniferales, Gnetales. Cambridge University Press.

Wagner, R. H. 2005. Dicranophyllum glabrum (Dawson) Stopes, an unusual element of lower Westphalian floras in Atlantic Canada. Revista Española de Paleontología 20 (1): 7-13.

The Sculpins of Baikal

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

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

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

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

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

Golomyanka Comephorus dybowskii, from here.

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

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


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

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

The Acrotretids: Micro-brachiopods from the Dawn of... Brachiopods

Ventral valve of Acrotreta sp., copyright Ivo Paalits / TÜ geoloogiamuuseum.

When brachiopods have been featured on this site before, they have generally been representatives of the group known as the articulates. Today's subjects, the Acrotretidae, are instead members of the inarticulate brachiopods. Whereas the shells of articulate brachiopods have a hinge connecting the two valves, the shells of inarticulates do not. Instead, the valves of inarticulates are held together purely by the muscle and tissue around them. Fewer of the living brachiopods are inarticulates than articulates, and the inarticulates have been less diverse over most of brachiopod history.

The Acrotretidae are one of the earliest known families of brachiopods in the fossil record, first appearing in the early Cambrian. They were most diverse in the later Cambrian and early Ordovician, becoming less so in the later Ordovician. Only a single genus is known to have survived into the Silurian (Holmer & Popov 2000). This may be something of a pseudo-extinction: the 'Acrotretidae' as currently defined is probably ancestral to other families of the order Acrotretida that post-dated it. Nevertheless, the acrotretid lineage as a whole became extinct during the Devonian. At one time it was thought that some living brachiopod families (the craniids and discinids) might be descendants of the acrotretids; they are now believed to not be closely related.

Reconstruction of the anatomy of the acrotretid Linnarssonia constans (with a boring parasite at lower left) from Bassett et al. (2004).

The first feature that springs to attention about the acrotretids is that they were tiny. In general, their shells were only one or two millimetres across. The two valves of the shell were generally quite distinct for each other. The dorsal valve was generally low and convex, whereas the ventral valve was more or less a deep lop-sided cone. A rounded or oval opening was present in the ventral valve, usually just behind the point of the cone. In life, this would have been the opening through which extended the pedicel, the fleshy stalk that would have attached the stalk to its substrate. In brachiopods as small as acrotretids, the lophophore would have been fairly simple. Living forms with such simple lophophores open the shell wide when feeding and hold the lophophore filaments in a bell-shape; water containing food particles is drawn into the centre of the 'bell' and pushed out laterally through the filaments (Rudwick 1965).

An alternate model of the acrotretid anatomy was proposed by Chuang in the early 1970s. He compared acrotretids to the living inarticulate brachiopod Lingula, in which the pedicel does not pass through an opening in the ventral valve but instead is positioned in the centre rear of the animal, passing between the two valves. Chuang suggested that the acrotretid pedicel did likewise, and that the opening in the conical valve (which he interpreted as dorsal rather than ventral) was used to expel water after it was drawn over the lophophore. In support of this model, he conducted an experiment in which he drilled holes in a comparable position in the dorsal valve of living craniid brachiopods (demonstrating once again the concept that one can get away with anything so long as one is experimenting on 'lower lifeforms'), through which the brachiopods did indeed expel water. However, Chuang's model was dismissed by Rowell (1977) who identified a number of features confirmed that the perforate valve of acrotretids was indeed ventral. Lingula, despite being the best-known inarticulate in the modern brachiopod fauna, is a poor model for acrotretids due to its adaptations to an infaunal lifestyle buried in mud, including the modification of the pedicel into a supersized structure for digging and anchoring itself. As for Chuang's experimental observations, Rowell argued that the only thing they demonstrated was that "a system under pressure leaks when perforated", noting that "This relationship... applies equally to bicycle tires and brachiopods".

So how did acrotretids make their living? The impression I've gotten while researching this post is that they are common in deposits that would have been part of the outer continental shelf. In particular, they are often found in black shales, a rock type that was originally formed from anoxic mud. Obviously, few animals are actually able to make a living in an environment lacking oxygen. Some do, such as the "rat-tailed maggot" larvae of hoverflies that possess a long breathing tube with which to obtain air, but it is difficult to imagine acrotretids functioning in this way. The other animals found fossilised in black shales alongside acrotretids are planktonic and nektonic forms, such as graptolites or cephalopods. It is possible that many acrotretids were pseudoplankton, living attached to other organisms or objects floating in the water, such as floating seaweeds (not floating wood, though, because wood didn't exist yet). When the acrotretid died, or its host substrate disintegrated, then it would begin the long descent towards eventual fossilisation in the black muds deep below.


Bassett, M. G., L. E. Popov & L. E. Holmer. 2004. The oldest-known metazoan parasite? Journal of Paleontology 78 (6): 1214–1216.

Holmer, L., & L. Popov. 2000. Lingulata. In: Kaesler, R. L. (ed.) Treatise on Invertebrate Paleontology pt H. Brachiopoda, Revised vol. 2. Linguliformea, Craniiformea and Rhynchonelliformea (part) pp. 30–146. Geological Society of America: Boulder, and University of Kansas: Lawrence.

Rowell, A. J. 1977. Valve orientation and functional morphology of the foramen of some siphonotretacean and acrotretacean brachiopods. Lethaia 10: 43-50.

Rudwick, M. J. S. 1965. Ecology and paleoecology. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt H. Brachiopoda vol. 1 pp. H199–H214. The Geological Society of America, Inc., and The University of Kansas Press.