Centaurea acaulis, Stemless Star-thistle

In an earlier post, I commented on the diversity of species of the star-thistle genus Centaurea. Among the many, many species that have been assigned to this genus is the stemless star-thistle Centaurea acaulis* of northern Africa.

*Though dissolution of the polyphyletic Centaurea may lead to this species changing places. Banfi et al. (2005) listed it under the name of Colymbada acaulis.

Patch of stemless star-thistles Centaurea acaulis, from L'herbiel de Gabriel.

Centaurea acaulis is an inhabitant of dry, rocky habitats that is native to Tunisia and northeastern Algeria. As indicated by both the vernacular and botanical names, its growth habit lacks a central stem. Instead, the long, lobed leaves (which can be up to about a foot in length going by photos provided by Agut Escrig et al., 2021) lie prostrate on the ground. These leaves end in a large, ovate apical section with lobes running down the side of the central rib, becoming smaller towards the base. Flower heads are solitary and carry a mass of bright yellow florets. The involucral bracts (the 'scales' around the outside of the base of the flower head) are flat and green with darker longitudinal veins. The distal section of the bracts is triangular with a membranous, ciliate margin and typically (though not always) ends in a long spine. A closely related species found in northwestern Algeria and Morocco, C. oranensis, has historically been treated as a subspecies of C. acaulis (under the name C. acaulis ssp. boissieri, because botanical nomenclature is weird). However, C. oranensis was raised to species level by Greuter & Aghababian (in Greuter & von Raab-Straube, 2005) on the basis of its distinct involucral bracts, which are distally blackish, ovate and concave, with a margin of dense, long, stiff setae.

Close-up of flower head of Centaurea acaulis, copyright Stephen Mifsud.

Recent years have seen this species extending its range northwards with populations now found in Spain, Italy and Malta. In Malta, it was initially found grown in a disturbed area with particularly alkaline soil (Buttigieg & Lanfranco 2001). The mechanism of its arrival is uncertain. It could have dispersed naturally across the Mediterranean, or it may have arrived mixed into bird seed. However it got there, one might expect that as the south of Europe becomes increasingly hotter and drier, the stemless star-thistle will continue to spread.

REFERENCES

Agut Escrig, A., J. P. Solís Parejo & P. Urrutia Uriarte. 2021. Noticias sobre la presencia de Centaurea acaulis L. (Asteraceae) en la Península Ibérica. Flora Montiberica 81: 51–54.

Banfi, E., G. Galasso & A. Soldano. 2005. Notes on systematics and taxonomy for the Italian vascular flora. 1. Atti Soc. It. Sci. Nat. Museo Civ. Stor. Nat. Milano 146 (2): 219–244.

Buttigieg, R., & E. Lanfranco. 2001. New records for the Maltese flora: Centaurea acaulis L. (family: Asteraceae). Central Mediterranean Naturalist 3 (3): 147–148.

Greuter, W., & E. von Raab-Straube (eds) 2005. Euro+Med notulae, 1. Willdenowia 35: 223–239.

Lifestyles of the Rosalinidae

Among the modern foraminiferans, one of the most prominent radiations is among members of the Rotaliida, characterised by globose chambers and calcareous, hyaline test walls. Among the numerous families making up the Rotaliida are members of the Rosalinidae.

Benthic form of Rosalina globularis, from Brady (1884).

Rosalinids may be regarded as fairly typical-looking marine rotaliids with the test growing freely as a low trochospire (so a flattened cone or dish shape). The aperture of the test is a low slit on the interior margin along the umbilicus (Hansen & Revets 1992). Rosalinids have a complex life cycle involving both benthic and planktonic stages (Sliter 1965). The asexually reproducing diploid stage is benthic. Depending on conditions, diploid individuals may divide to produce other diploid individuals, resulting in several asexual generations. Eventually, however, the diploid generation will undergo meiosis to produce the haploid sexual generation (in the common species Rosalina globularis, this is induced by exposure to warmer water). In the sexual generation, a large globular chamber forms at maturity that covers the umbilical side of the test. This float chamber becomes filled with gas, allowing the foram to disperse planktonically before releasing gametes to produce the next diploid generation. Planktonic individuals are distinct enough in appearance from their benthic counterparts that they were long mistaken for distinct taxa before their identity was revealed by lab cultures.

Life cycle of Rosalina globularis, from Sliter (1965).

The majority of forams are particulate feeders. A network of filamentous pseudopodia radiating outwards from the cell body captures micro-organisms and other organic particles. However, one genus of rosalinids, Hyrrokkin, lives as parasites on sessile invertebrates (Cedhagen 1994). Species of this genus have variously been found on sponges, corals and bivalves. On sponges, they settle on the inhalent surface of the sponge and dissolve the underlying tissues. On bivalves, they form pits on the shell surface from which they bore holes through to the body cavity. Pseudopodia extended through this hole allow the foram to feed on host tissue. Infested hosts may bear multiple scars from the foram moving about on the outer surface. The forams may also feed on other animals such as polychaete worms or bryozoans attached to the surface of their primary host. In such cases, Hyrrokkin remains in its original pit but develops an irregularly shaped chamber with its aperture directed towards the alternate prey. Hyrrokkin species evidently do well from their rapacious lifestyle: whereas other rosalinids are only a fraction of a millimetre in diameter, Hyrrokkin sarcophaga is an absolute giant reaching around six millimetres across and with protoplasm containing thousands of nuclei. Proving once again that one may make a great deal of profit from the labour of others.

Cross-section of Hyrrokkin sarcophaga boring into shell of file clam Acesta excavata, from Schleinkofer et al. (2021).

REFERENCES

Cedhagen, T. 1994. Taxonomy and biology of Hyrrokkin sarcophaga gen. et sp. n., a parasitic foraminiferan (Rosalinidae). Sarsia 79: 65–82.

Hansen, H. J., & S. A. Revets. 1992. A revision and reclassification of the Discorbidae, Rosalinidae, and Rotaliidae. Journal of Foraminiferal Research 22 (2): 166–180.

Sliter, W. V. 1965. Laboratory experiments on the life cycle and ecologic controls of Rosalina globularis d'Orbigny. Journal of Protozoology 12 (2): 210–215.

Colus and Co.

The neogastropods have long been a challenge taxonomically. They are extremely diverse, encompassing a large number of species with a wide range of lifestyles, but they also exhibit exhibit regular patterns of convergence and/or conservatism between different lineages. Perhaps the most challenging group of all has been the whelks, commonly recognised as the superfamily Buccinoidea, a massive radiation of over 3300 known species. Whelks are particularly diverse in colder regions of the world's oceans, including amongst their number there the members of the family Colidae.

Hairy colus Colus pubescens, copyright E. A. Lazo-Wasem.

Colus has been used as the basis of a family group name at many levels of whelk classification, whether it be Colidae, Colinae or Colini. The gastropod classification laid out by Bouchet et al. (2017) recognised 'Colini' as a diverse tribe within the main whelk family Buccinidae, including a range of cold-water taxa. However, a more recent phylogenetic analysis of the buccinoids by Kantor et al. (2021) found Bouchet et al.'s concept of Colini to be polyphyletic, placing the type genus Colus outside what the called the 'core Buccinoidea'. As such, they raised Colidae to the status of a separate family and restricted it to just two genera, Colus and Turrisipho.

In this restricted form, the Colidae are thin-shelled, medium-sized to large whelks with the largest having shells up to twenty centimetres in length. The shells are fusiform to ovate in shape with a more or less elongate siphonal canal and covered by a brown periostracum. Axial sculpture is absent; spiral sculpture is expressed as more or less prominent cords. The aperture is closed with a operculum bearing a terminal nucleus. The animal has a more or less long proboscis. The radula bears three teeth per row; the middle tooth has a more or less square base and one to three cusps, with the middle cusp the largest, whereas the lateral teeth bear three hooked cusps with the outermost cusp significantly larger than the other two. None of these features, it should be noted, is entirely unique to the Colidae (Kantor et al. 2021).

Turrisipho dalli, from BoldSystems.

Members of the Colidae are found in the Arctic and northern Atlantic Oceans, from subtidal to bathyal depths. Because they are not targeted commercially, the life habits of colids have not been well studied. However, what we do know indicates that they are likely predators on other invertebrates (Kosyan 2007). The long proboscis of most species is probably used to pull infaunal animals such as amphipods and bivalves out of their burrows. Colids have well-developed salivary glands and it is possible that these may produce toxins as found in other neogastropods. They do not have anything like the elaborate venom delivery setups like those found in the conoids, but even a little dose of toxic saliva helps to subdue a struggling crustacean.

REFERENCES

Bouchet, P., J.-P. Rocroi, B. Hausdorf, A. Kaim, Y. Kano, A. Nützel, P. Parkhaev, M. Schrödl & E. E. Strong. 2017. Revised classification, nomenclator and typification of gastropod and monoplacophoran families. Malacologia 61 (1–2): 1–526.

Kantor, Y. I., A. E. Fedosov, A. R. Kosyan, N. Puillandre, P. A. Sorokin, Y. Kano, R. Clark & P. Bouchet. In press 2021. Molecular phylogeny and revised classification of the Buccinoidea (Neogastropoda). Zoological Journal of the Linnean Society.

Kosyan, A. R. 2007. Morphological features, ecology, and distribution of poorly studied molluscan genera of the Colinae subfamily (Gastropoda, Buccinidae) from the far eastern seas of Russia. Oceanology 47 (4): 531–536.