Bucklandiella lusitanica

The diversity of mosses is much higher than many people realise. Whereas some moss species have wide ranges that may cross between continents and hemispheres, others are unique to very specific regions and habitats. Among examples of the latter is the European species Bucklandiella lusitanica.

Illustrations of Bucklandiella lusitanica, from Ochyra & Sérgio (1992). Top left: habit; top right: section of stem of hair-leafed form when dry; lower left: section of stem of hairless form and sporophyte when wet.

Bucklandiella lusitanica was only described as a new species (under the name Racomitrium lusitanicum) in 1992 (Ochyra & Sérgio 1992), having gone unnoticed previously despite being a relatively distinctive species. Recent collections of the species have been identified from a single region, the Serra do Gerês mountain rainge and Parque Natural da Peneda-Gerês national park in the northwest of Portugal, at altitudes between 650 and 1000 metres. A single collection from the Serra do Estrela further south in the country was made in the mid-1800s though it went unidentified at the time. Its rarity is such that is has officially been listed as Endangered by the IUCN. Bucklandiella lusitanica is a rheophyte, which is to say that it grows in association with running water. It grows on acidic granite rocks that are periodically or permanently submerged, such as alongside streams and waterfalls. It is particularly abundant on steep rock faces, growing in association with closely related moss species.

Appearance-wise, Bucklandiella lusitanica is a medium-sized moss with irregularly branched stems growing 1.5 to 3.5 centimetres in length. Leaves are rigid, held tight to stem, and two or three millimetres long.One of the species' most distinctive features is a broad, fleshy margin to each leaf that is generally two or three cells thick whereas the lamina of the leaf is mostly only a single cell thick. The alar cells at the base of the sides of the leaf often form inflated, strongly coloured lobes. The leaves commonly end in a fine, colourless hair-point. The structure of the leaves is similar to that of Bucklandiella lamprocarpa, another aquatic moss species, but that species lacks the hair-points. The two species also differ in the form of their spores, those of B. lamprocarpa being larger and more ornate than those of B. lusitanica, and B. lamprocarpa has fatter and often shinier capsules than B. lusitanica.

I mentioned previously that Bucklandiella lusitanica was originally described as a member of the genus Racomitrium. The moss genus Racomitrium was long recognised by a distinctive array of features including leaf lamina cells with distinctly sinuous longitudinal cell walls, a calyptra (the cap of the developing capsule) that is basally frayed into several lobes, and teeth of the peristome (the teeth around the aperture of a mature capsule) that are split into two or more segments (Sawicki et al. 2015). Racomitrium in this sense was a diverse genus with over two hundred species having been named at one time or another, and somewhere between sixty and eighty species recognised as valid in recent years, As a result, Ochyra et al. (2003) proposed the division of Racomitrium in the broad sense between four separate genera. Bucklandiella, the largest of these segregate genera (with about fifty currently known species), was recognised for species with a smooth leaf surface (lacking papillae on the lamina) and relatively short, shallowly divided teeth in the peristome. The division of Racomitrium has not been universally accepted. Larrain et al. (2013) questioned the monophyly and diagnosability of Ochyra et al.'s segregates but Sawicki et al. (2015) reiterated their support for the new system (and added a fifh new segregate genus to boot). It is generally accepted that Racomitrium in the broad sense represents a monophyletic unit, so the question of whether lusitanicum should be assigned to Racomitrium or Bucklandiella may largely be considered a question of just how closely circumscribed you feel a genus should be.

REFERENCES

Larraín, J., D. Quandt, M. Stech & J. Muñoz. 2013. Lumping or splitting? The case of Racomitrium (Bryophytina: Grimmiaceae). Taxon 62 (6): 1117–1132.

Ochyra, R., & C. Sérgio. 1992. Racomitrium lusitanicum (Musci, Grimmiaceae), a new species from Europe. Fragmenta Floristica et Geobotanica 37 (1): 261–271.

Ochyra, R., J. Żarnowiec & H. Bednarek-Ochyra. 2003. Census Catalogue of Polish Mosses. Institute of Botany, Polish Academy of Sciences: Cracow.

Sawicki, J., M. Szczecińska, H. Bednarek-Ochyra & R. Ochyra. 2015. Mitochondrial phylogenomics supports splitting the traditionally conceived genus Racomitrium (Bryophyta: Grimmiaceae). Nova Hedwigia 100 (3–4): 293–317.

The Glandulinid Position

In an earlier post, I described how the majority of modern multi-chambered foraminiferans can be divided between two lineages, the Tubothalamea and Globothalamea. The two groups generally differ in the shape of the first chamber following the proloculus (the central embryonic chamber of the test): in one, this chamber is tubular whereas in the other it is globular or crescent-shaped (guess which is which). But there is a third notable group of multi-chambered forams: the Nodosariata. In both tubothalameans and globothalameans, the chambers more or less coil around the proloculus to form a spiral. In the Nodosariata, the test is more or less linear with apical chamber apertures. The chambers may be successively stacked one after the other to form a uniserial test, or they may be arranged in a zig-zag or twirling arrangement to form biserial, triserial, etc. arrangments. In living Nodosariata, the wall of the test is made of a single layer of hyaline calcite though some earlier representatives (up to the end of the Jurassic) had differing wall make-ups (Rigaud et al. 2016). Among the numerous notable representatives of the Nodosariata in the modern fauna are representatives of the family Glandulinidae.

Series of Glandulina ovula, from Brady (1884).

Species have been assigned to the Glandulinidae going back to the Jurassic with the modern genus Glandulina recognisable in the Palaeocene (Loeblich & Tappan 1964). The test may be uniserial, biserial or polymorphine (more than two series); a common arrangement is for the test to start out biserial or polymorphine then become uniserial as the individual chambers become larger. In Glandulina, the microspheric generation starts biserial but the megalospheric form is uniserial throughout (Taylor et al. 1985). As the test grows, the internal walls between chambers may be resorbed. The terminal aperture of the test may be radial or slit-like. The most characteristic feature of the family is a tube running into the chamber from the inside of the aperture, referred to as the entosolenian tube. Some glandulinids have been described as lacking an entosolenian tube but such absences are likely artefacts of preservation: the delicate tube is easily dislodged during the fossilisation process (Taylor et al. 1985).

The overall relationships of the Nodosariata remain a question open to investigation. The classification of forams by Loeblich & Tappan (1964) included both multi-chambered and single-chambered (unilocular) forms within the Glandulinidae, with the unilocular forms placed in a subfamily Oolininae. Oolinines resemble glandulinids proper in a number of features including wall structure and the presence of an entosolenian tube. More recent authors, however, have rejected this relationship. Rigaud et al. (2016) entirely excluded unilocular forms from the Nodosariata as a whole, regarding it as improbable that single-chambered forms could have evolved from multi-chambered ancestors (as would seemingly be required by their relative appearances in the fossil record). Do the similarities between glandulinids and oolinines reflect a common ancestry, or are they the result of simple convergence? Unfortunately, with so few significant characters available to inform our understanding of foram higher relationships, the answer you prefer may come down to no more than your own personal feelings about which indicators are more reliable.

REFERENCES

Loeblich, A. R., Jr, & H. Tappan. 1964. Treatise on Invertebrate Paleontology pt C. Protista 2. Sarcodina: chiefly "thecamoebians" and Foraminiferida vol. 2. The Geological Society of America, and The University of Kansas Press.

Rigaud, S., D. Vachard, F. Schlagintweit & R. Martini. 2016. New lineage of Triassic aragonitic Foraminifera and reassessment of the class Nodosariata. Journal of Systematic Palaeontology 14 (11): 919–938.

Taylor, S. H., R. T. Patterson & H.-W. Choi. 1985. Occurrence and reliability of internal morphologic features in some Glandulinidae (Foraminiferida). Journal of Foraminiferal Research 15 (1): 18–23.

Key Limpets

On two occasions before, I've presented you with members of the Fissurellidae, the keyhole and slit limpets. It's time for a return visit to the fissurellids, in the form of the diverse keyhole limpet genus Diodora.

Various views of shell of Diodora italica, copyright H. Zell.

Species have been assigned to Diodora from coastal waters pretty much around the world except for the coolest regions. They are small to moderate-size limpets, the largest species growing about three centimetres in length and two centimetres in height. The shell opens through a 'keyhole' at the apex through which the animal ejects waste matter and water that has been passed over the gills. The internal margin of this keyhole is surrounded by a callus on the underside of the shell; a distinguishing feature of Diodora is that this callus is posteriorly truncate. The external ornament of the shell is cancellate (arranged in a criss-cross pattern) and the margin of the shell is internally crenulated (Moore 1960). Moore (1960) listed three subgenera of Diodora distinguished by features of the keyhole shape and position but Herbert (1989) notes that these subgenera are not clearly distinct. A phylogenetic analysis of the fissurellids by Cunha et al. (2019) did recognise a clade including the majority of Diodora species analysed. However, species from the eastern Pacific formed a disjunct clade that may prove to warrant recognition as a separate genus.

As far as is known, Diodora species have a long lifespan, surviving for some ten to twenty years. They do not have a planktonic larva; young Diodora hatch directly from the egg as benthic crawlers. For the most part, they are presumed to graze on algae in the manner of other fissurellids and limpets. However, the northeastern Australian species D. galeata has been found feeding on the soft tissues of coral (Stella 2012), a habit that went unrecognised until fairly recently owing to the animal's cryptic nature, hiding deep among the branches of the host. Whether other Diodora species might exhibit similar lifestyles would require further investigation.

REFERENCES

Cunha, T. J., S. Lemer, P. Bouchet, Y. Kano & G. Giribet. 2019. Putting keyhole limpets on the map: phylogeny and biogeography of the globally distributed marine family Fissurellidae (Vetigastropoda, Mollusca). Molecular Phylogenetics and Evolution 135: 249–269.

Herbert, D. G. 1989. A remarkable new species of Diodora/i> Gray, 1821 from south-east Africa (Mollusca: Gastropoda: Fissurellidae). Annals of the Natal Museum 30: 173–176.

Moore, R. C. (ed.) 1960. Treatise on Invertebrate Paleontology pt I. Mollusca 1. Mollusca—general features, Scaphopoda, Amphineura, Monoplacophora, Gastropoda—general features, Archaeogastropoda and some (mainly Paleozoic) Caenogastropoda and Opisthobranchia. Geological Society of America, Inc. and University of Kansas Press.

Stella, J. S. 2012. Evidence of corallivory by the keyhole limpet Diodora galeata. Coral Reefs 31: 579.

Small Carpenters

It's time for another dip into the wide diversity of bees. The small carpenter bees of the tribe Ceratinini are small (often less than a centimetre in length), slender bees found on all continents except Antarctica, though their toehold in Australia is a very tenuous one indeed with only a single known species there. Though diverse, with hundreds of known species, the difficulty of breaking the tribe into clearly defined, monophyletic groups has lead recent authors to recognise a single genus Ceratina (Michener 2007). Distinctive subgroups previously treated as separate genera, such as the relatively large Megaceratina and the heavily punctate Ctenoceratina of Africa, and the both bright metallic and heavily punctate Pithitis of Africa and southern and eastern Asia, are now treated as subgenera. There are a lot of recognised subgenera, over twenty at last count, but there are also a lot of species not yet assigned to subgenus. Phylogenetic analysis of the ceratinins supports monophyly of most subgenera and a likely African origin for the clade as a whole, with multiple dispersals into Eurasia followed by a single dispersal to the Americas (Rehan & Schwarz 2015).

Ceratina sp., possibly C. smaragdula, copyright Vengolis.

Distinctive features of Ceratina compared to other bees include the absence of a pygidial plate, a flattened and hardened patch on the tip of the abdomen in females. As members of the family Apidae, Ceratina are long-tongued bees with a scopa (cluster of pollen-carrying hairs) on the hind legs, though the scopa does not enclose a bare patch for carrying a shaped pollen ball as in some other apids (for instance, the familiar honey bees). The scopa is less extensive in small carpenter bees than it is in other apids and the hairs on the body as a whole are rather short, so Ceratina look much shinier and less fuzzy than other bees. Ceratina are black or metallic green in colour (on rare occasions, the metasoma is red) and usually have yellow patches, particularly on the face.

Ceratina nest in a fennel stem, copyright Gideon Pisanty.

The name 'carpenter bee' refers to their practice of nesting in hollow stems or twigs, entered at broken ends. The absence of the pygidial plate is probably related to this manner of nesting: it is normally used by ground-nesting bees to tamp down soil when closing the nest. Most of the time, Ceratina are solitary nesters but two or more females may sometimes work on a nest together. In these cases, they adopt a proto-eusocial division of labour with one female laying eggs while the others act as 'workers' (I have no idea how they decide who gets to do what). Though a reduction in hairiness in bees is often associated with kleptoparasitism, no Ceratina species are known to behave in that manner (though some kleptoparasites are known among the members of the closely related and very similar tribe Allodapini). The reduction of the scopa may instead be associated with the bees carrying food supplies for the nest in their crop as well as on the legs. Cells are lined up in the nest stem with only simple partitions between them. These partitions are made from loose particles, mostly the pith of the stem, with no obvious adhesive holding them together. In at least some species, females will return to the nest after completion, dissembling and reassembling cell walls in order to clean out dead larvae and faeces that are then incorporated into the partitions. As such, while small carpenter bees are not directly on the evolutionary line leading to the more integrated colonies of the social bees, they do provide us with a model of what one stage in their evolution may have looked like.

REFERENCES

Michener, C. D. 2007. The Bees of the World 2^nd ed. John Hopkins University Press: Baltimore.

Rehan, S., & M. Schwarz. 2015. A few steps forward and no steps back: long-distance dispersal patterns in small carpenter bees suggest major barriers to back-dispersal. Journal of Biogeography 42: 485–494.

Rove by the Riverside

The Staphylinidae, commonly known as the rove beetles, are one of the most diverse of the recognised beetle families. Indeed, thanks to their habit in recent years of glomming up lineages previously treated as distinct families like the pselaphids and scydmaenids, they now rival the weevils of the Curculionidae for the position of largest of all recognised animal families. But for their diversity and ubiquity, staphylinids are comparatively poorly studied, owing to a not-unwarranted reputation for taxonomic recalcitrance (the relatively soft bodies of many staphylinids mean they often do not handle well with standard methods for examining beetles). Perhaps the most neglected of all staphylinid subgroups is the subfamily Aleocharinae. Aleocharines are often minute (the average aleocharine is only a couple of millimetres in length) and their identification often requires resolving features that lie at the very limit of what can be seen with a standard dissecting microscope. Nevertheless, aleocharines are remarkably diverse and among their representatives are the representatives of the genus Parocyusa.

Parocyusa americana, from Brunke et al. (2012); scale bar = 1 mm.

Typical aleocharines have what is thought of as the 'standard' body form for staphylinids, with short, square elytra that do not cover the long, flexible abdomen (though I should mention that, with the aforementioned assimilation of the pselaphids and scydmaenids, I suspect there may now be more 'non-standard' staphylinid species than 'standard' ones). For the most part, they can be distinguished from other staphylinid subfamilies by the position of the antennae, with their insertions placed behind the level of the front of the eyes. Aleocharines are divided between numerous tribes; Parocyusa is included in the tribe Oxypodini, a heterogenous group of relatively unspecialised aleocharines. Notable features distinguishing Parocyusa from other aleocharine genera include legs with five segments to each tarsus, a frontal suture between the antennal insertions, the median segments of the antennae being longer than wide, the head not having a well defined 'neck', the sides of the pronotum not being strongly deflexed downwards (so the hypomeron, the lateral section of the pronotum, is clearly visible in side view), and deep transverse impressions across the third to fifth abdominal tergite but not across the sixth tergite or across the sternites (Newton et al. 2001). Members of the genus are a bit over three millimetres in length.

Species of Parocyusa are found widely in the Holarctic realm; I've found reference to species from Europe, Korea, and northeastern North America (I should also note that I've also encountered dark allusions to recent rearrangements of the generic status of some of these species but without access to such revisions I'm going to stick with what I can find). I haven't found any reference to their specific diet but I suspect that they would be micropredators, a common lifestyle for staphylinids of their kind. Parocyusa species are associated with running water, living among the gravel and sand alongside stream beds (e.g. Brunke et al. 2012). As such, these and other aleocharines have received attention in ecological studies: the higher the diversity of staphylinids present, the more healthy the ecosystem is likely to be.

REFERENCES

Brunke, A. J., J. Klimaszewski, J.-A. Dorval, C. Bourdon, S. M. Paiero & S. A. Marshall. 2012. New species and distributional records of Aleocharinae (Coleoptera, Staphylinidae) from Ontario, Canada, with a checklist of recorded species. ZooKeys 186: 119–206.

Newton, A. F., M. K. Thayer, J. S. Ashe & D. S. Chandler. 2001. Staphylinidae Latreille, 1802. In: Arnett, R. H., Jr & M. C. Thomas (eds) American Beetles vol. 1. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia pp. 272–418. CRC Press: Boca Raton.

Caloppiidae

The concept of ranks in taxonomy is ultimately an arbitrary one. There is no real definition of what constitutes an 'order', a 'family' or a 'subfamily'. What determines the rank that a given taxon is recognised at is a combination of tradition, convenience, and the taxon's relationships to other recognised taxa. As such, the question of whether a given classification is overly 'split' or 'lumped' is a meaningless one and arguing the point is a complete waste of time. That said, the classification of the 'higher' oribatid mites is massively oversplit.

A big part of the reason why oribatid classification seems such a mess, with large numbers of small families containing only a handful of genera and/or species apiece, can be attributed to simple ignorance. We simply do not have a good handle on how many oribatid taxa are related to each other and as a result we find ourselves with a great many orphan taxa still hunting for a good home. The Caloppiidae may be regarded as one such taxon.

Dorsal view of Luissubiasia microporosa, from Ermilov (2016). Scale bar = 100 µm; labels with 'A' indicate areae porosae.

Caloppiids are a pantropical group of about thirty species of poronotic oribatids (the group of oribatids exhibiting the octotaxic system, an arrangement of glandular openings on the notogaster), with three genera recognised in the family by Ermilov (2016): Zetorchella, Brassiella and Luissubiasia. Zetorchella, which includes the majority of the family's species, is also pantropical in distribution. Brassiella is known from the Indo-Pacific region and Liussubiasia is known from a single species from Cuba. Past authors have often referred to Zetorchella and the Caloppiidae by the names Chaunoproctus and Chaunoproctidae, respectively, but as the name Chaunoproctus had already had dibs called on it before the mite was named (by a bird, the now-extinct Bonin grosbeak Chaunoproctus ferreorostris), their respective most senior synonyms have to take over. Caloppiids are more or less egg-shaped in dorsal view. They lack the distinct pteromorphs of most other poronotics though they may have quadrangular projections in the humeral region (the 'shoulders'). The integument is usually heavily sculpted and foveate. The legs end in three claws apiece. The most characteristic feature of the group is that the openings of the octotaxic system on the notogaster, of which five pairs are present, are extremely small. The octotaxic system can take two forms, recessed saccules or porose patches. Those of caloppiids have usually been described as saccules but Ermilov (2016) states that, at least in some species, they are very small porose areas.

Going by their overall appearance, caloppiids are classified within the superfamily Oripodoidea. However, one of the most characteristic features of the Oripodoidea as an evolutionary group is that their nymphs have notogastral setae borne on individual off-centred sclerites (oribatid nymphs often look very different from their adults and are often more soft-bodied). At this point in time, we simply do not know what the nymphs of caloppiids look like so we cannot say whether they possess this crucial feature. Conversely, with their lack of pteromorphs, caloppiids bear a distinct similarity to the more diverse oripodoid family Oribatulidae. The two families have mostly been separated on the basis of caloppiids supposedly having an octotaxic system of saccules rather than porose areas, a distinction that I've already noted may not hold up. There's also something of an open question whether the distinction between saccules and porose areas is really as significant as it has been thought in the past. So, at present, we can't say with confidence whether caloppiids are true oripodoids... or whether they are not only oripodoids but don't even warrant recognition as a distinct family from oribatulids.

REFERENCE

Ermilov, S. G. 2016. Luissubiasia microporosa gen. nov., sp. nov. (Acari, Oribatida, Caloppiidae) from Cuba. International Journal of Acarology 42 (2): 127–134.

The Grisons

Spend a bit of time following discussions of nature documentaries and other popular representations of biodiversity, and one topic you're likely to see come up is the biases that tend to exist in what gets represented. Images from eastern and southern Africa predominate while the west and north of that continent get overlooked. Europe and North America receive much more attention than the temperate regions of Asia. Another region whose diversity tends to go underrepresented is South America. The casual observer might think this continent is all monkeys and jaguars but South America is also home to notable radiations of dogs, deer, rodents, and other animals that many people would associate more with other parts of the world. Among these overlooked elements of the South American fauna are the local species of mustelid, including the grisons of the genus Galictis.

Greater grison Galictis vittata, copyright Tony Hisgett.

Grisons are somewhat ferret- or skunk-like animals found across almost the entirety of South America, and north into southern Mexico. They are greyish in colour dorsally (the name 'grison' itself means 'grey') with a black face and underparts. A pale stripe separates the upper and lower parts across the top of the face and continues diagonally back to the shoulders. They feed on small vertebrates and tend to be solitary hunters though they may sometimes form small family groups. They are primarily terrestrial and diurnal in habits. They have a reputation for ferocity; residents of Chile apparently have a history of using comparisons to grisons to describe unchecked rage (Yensen & Tarifa 2003b), in a similar manner to references to wolverines and honey badgers in other parts of the world. Contrasting colour patterns like those of the grisons are associated in other musteloids (such as skunks) with the production of offensive odours for defence, and grisons also produce strong-smelling secretions from their anal glands. Though some sources have claimed the odour produced by the lesser grison to be worse than a skunk's, it appears that these reports are exaggerated (Yensen & Tarifa 2003b).

Lesser grison Galictis cuja, copyright Ken Erickson.

Most authors have recognised two species of grison, the greater grison Galictis vittata and the lesser grison G. cuja*, as corroborated by a recent taxonomic study of the genus by Bornholdt et al. (2013). As their names indicate, the greater grison is generally larger and more robust than the lesser, being about 60 to 76 cm in total length versus 44 to 68 cm for the lesser grison (Yensen & Tarifa 2003b). The tail is also proportionately shorter in the greater grison (30% of the total length for the greater, 40% for the shorter). Fur is relatively longer and denser in the lesser grison, giving it more of a fluffy look. Whereas the dorsal fur is always a plain grey in the greater grison, it may often have a yellowish tinge in the lesser (not always, though). The two are generally distinct in range and habitat, as well. The greater grison is an animal of tropical forests and inhabits the northern part of the genus' range in Central America and northern and western South America. The lesser grison inhabits drier habitats, in arid or temperate regions, and so occupies the southern and eastern parts of the continent. The ranges of the species are known to overlap in Bolivian and Paraguay where their respective biomes approach each other.

*Some sources have listed a third species G. allamandi but this seems have been something of a 'ghost' taxon born from confusion whether the name 'G. vittata' applied to the greater or lesser species.

The genus Galictis arrived in South America as part of the Great American Biotic Interchange, about three million years ago. The general consensus is that it is derived from the genus Trigonictis of the North American Pliocene. Indeed, it has even been suggested that the two North American species of Trigonictis might represent independent ancestors of Galictis, with the larger T. macrodon giving rise to the greater grison and the smaller T. cookii birthing the lesser grison (Yensen & Tarifa 2003a). This certainly would seem overly complicated, though, and molecular data are more in line with a more recent separation of the species.

REFERENCES

Bornholdt, R., K. Helgen, K.-P. Koepfli, L. Oliveira, M. Lucherini & E. Eizirik. 2013. Taxonomic revision of the genus Galictis (Carnivora: Mustelidae): species delimitation, morphological diagnosis, and refined mapping of geographical distribution. Zoological Journal of the Linnean Society 167: 449–472.

Yensen, E., & T. Tarifa. 2003a. Galictis vittata. Mammalian Species 727: 1–8.

Yensen, E., & T. Tarifa. 2003b. Galictis cuja. Mammalian Species 728: 1–8.