Field of Science

Meandering Forams

Specimen of Meandropsina vidali, showing the patterning on the external surface, from Loeblich & Tappan (1964).

There are some taxonomic names that just instantly bring up a mental image of the sort of organism to which they refer. For my part, I've always felt that Meandropsina is one of those names. The Meandropsinidae are another family of relatively large and complex foraminifera (growing up to a number of millimetres across) that are known only from the Upper Cretaceous. The several genera of the family are predominantly European, with only the genus Fallotia also known from the West Indies.

Cross-section of Meandropsina vidali, from Loeblich & Tappan (1964).

Meandropsinids are (as far as I know) more or less lenticular in shape with chambers enrolled in a flat spiral. The name of the type genus Meandropsina refers to the way that the outer margins of the chambers tend to meander irregularly around the test, giving it something of an ornate appearance. Both molecular and structural evidence indicate that multi-chambered forams arose from ancestors with undivided tests on more than one occasion, and the majority of multi-chambered forams can be assigned to two major lineages (Pawlowski et al. 2013). In one lineage, the Globothalamea (which includes, for instance, the rotaliids), the basic chamber shape is globular with successive chambers in the test being wider than long. In the other lineage, the Tubothalamea (including the miliolids and spirillinids), the basic chamber shape is tubular, and the test may grow through a number of spirals before it even starts to be divided into chambers (if at all). Members of the two lineages with calcareous tests may also be distinguished by their test structure: in calcareous globothalameans, the crystals making up the test are arranged regularly so the overall appearance of the test is hyaline (glass-like). In contrast, tubothalameans have the crystals of the test arranged irregularly so the appearance of the test is porcelaneous (like porcelain). Meandropsinids are unmistakeably tubothalameans in both regards.

Like other large forams of the Mesozoic, meandropsinids did not make it past the end of the Cretaceous. Early Palaeocene taxa that have been included in the families represent distinct lineages that evolved to take their place, occupying the ecological spaces opened up by the mass extinction ending the era.


Loeblich, A. R., Jr, & H. Tappan. 1964. Treatise on Invertebrate Paleontology pt C. Protista 2. Sarcodina: chiefly "thecamoebians" and Foraminiferida vol. 1. The Geological Society of America, and The University of Kansas Press.

Pawlowski, J., M. Holzmann & J. Tyszka. 2013. New supraordinal classification of Foraminifera: molecules meet morphology. Marine Micropalaeontology 100: 1–10.

Doe, it's Deer

Marsh deer Blastocerus dichotomus, copyright Jonathan Wilkins. An animal that just screams out, "Am I wearing the Chanel boots? Yes, I am."

I hardly need to explain what deer are, do I? Deers (Cervidae) are generally recognised as the second most diverse family of hoofed mammals (after bovids) in the modern fauna. Their most recognisable feature, of course, is the possession of antlers: bony cranial appendages that are shed and regrown every year rather than being permanently in place like the horns of a bovid. When antlers first grow, they are covered with a layer of skin (the velvet) that supplies them with blood, but this skin is later shed to expose the bare bone. In most species, antlers are only grown by males whose use them in conflicts during the mating season. The only genus of deer that grows antlers in both sexes is Rangifer, the reindeer. There is also one living species that lacks antlers, the Chinese water deer Hydropotes inermis; instead of antlers, males of this species possess large, dagger-like canines. In the majority of deer species, antlers are subcylindrical and often branched but broad palmate antlers have evolved on multiple occasions within the family. Antler morphology is generally significant in distinguishing taxa but it should be noted that variation within species is not unknown. For instance, the few recorded males of the small, now possibly extinct population of moose introduced to the south of New Zealand lacked the large palmate antlers generally associated with the species, probably due to poor nutritional conditions. Instead, they had more slender antlers that only became moderately palmate distally, like those of a fallow deer Dama dama.

One of the few photographs of moose from New Zealand, from here. I think this might be the one shot at Herrick Creek in 1952 but I could be wrong.

The first antlered deer are known from Europe back in the early Miocene, about 17 million years ago. They are not known from North America until some time later in the Pliocene, about five mya (Pitra et al. 2004), though these days they are every bit as diverse in the Americas as in Eurasia. They never made much inroad into Africa, only extending into the northernmost part of the continent, and they never made it into Australasia under their own steam, though a number of species have been dispersed to various parts of the world by humans. For instance, at least half a dozen species have become established in New Zealand, and until recently reindeer might be found wandering among penguin colonies in South Georgia.

Reindeer and king penguins on South Georgia, from here.

Recent decades have seen some pretty wild swings in cervid taxonomy, with the number of subfamilies recognised varying from two to seven, and some authors recognising a much larger number of genera and species than others. However, our general understanding of cervid interrelationships is pretty good these days, with many differences between systems being a question of ranking more than anything else. Recent studies have agreed that modern deer can be divided between two primary lineages that may be called the Cervinae and Capreolinae (Gilbert et al. 2006). The Cervinae include the majority of deer species in the Old World with a single species (the wapiti Cervus canadensis) extending its range into the New World. The remaining New World deer all belong to the Capreolinae, which also includes four genera (Rangifer, Hydropotes, the roe deer Capreolus and the moose Alces) found in Eurasia.

Male tufted deer Elaphodus cephalophus, copyright Heush.

The Cervinae can be divided between two tribes, the Muntiacini and Cervini. Muntiacini include the muntjaks of the genus Muntiacus and the tufted deer Elaphodus cephalophus. These are small deer native to southern and eastern Asia. Antlers are small and simple in all Muntiacini: muntjaks have antlers with only a single short anterior branch whereas the tufted deer has unbranched antlers that are barely visible under the large tuft of hair that this species has on top of the head. Muntiacini also resemble Hydropotes in their possession of large canines in the males. The other tribe, Cervini, includes larger deer species with more complex, multi-branched antlers. Some authors have historically placed all species of Cervini within a single genus Cervus; others may recognise nine distinct genera. Numbers of recognised species have also varied, largely due to phylogenetic studies finding that taxa previously recognised as conspecific subspecies may be more distantly related to each other or may not form monophyletic units. For instance, the wapiti has often been regarded as a subspecies of the red deer Cervus elaphus but recent studies have suggested that it is more closely related to the sika C. nippon and the white-lipped deer Przewalskium or Cervus albirostris, two east Asian species (Pitra et al. 2004). Difficulties in elucidating cervin phylogeny are probably best exemplified by the case of Père David's deer Elaphurus or Cervus davidianus, originally native to southern China but now only surviving in captivity. Molecular phylogenies associate this species closely with the brow-antlered deer Cervus eldi but it has many morphological features indicating a close relationship with C. elaphus, and it is widely suspected that Père David's deer originated from a hybridisation event between the two latter species.

Pudu (I think a northern pudu Pudu puda), copyright Neil McIntosh.

The Capreolinae can be divided between three main lineages. One comprises the roe deer and water deer; another comprises the moose (again, I'm making a point of referring to genera rather than species because the number of recognised species may differ between authors). Note that the position of the water deer suggests that the antler-less state of this species represents a secondary loss rather than retention of a primitive state. The majority of capreolines belong to the third lineage, commonly recognised as the tribe Odocoileini. Except for the reindeer, the species of this lineage are restricted to the New World, with the higher diversity in South America. The Odocoileini are perhaps the most taxonomically uncertain section of the deer family. There appears to be no question, at least, that Rangifer represents the sister group of all other Odocoileini. The remaining odocoileins have generally been divided between six genera: Odocoileus (including the mule deer O. hemionus and white-tailed deer O. virginianus), Mazama (brockets), Pudu (pudus), Hippocamelus (guemuls), the marsh deer Blastocerus dichotomus and the pampas deer Ozotoceros bezoarticus. However, except for the two monotypic genera, monophyly of all these taxa was placed in question by a recent molecular phylogenetic study of the group by Gutiérrez et al. (2017). The issue is particularly marked for the brockets, small deer with unbranched antlers, as not only the genus as a whole but also species within the genus have been indicated as non-monophyletic. Recent years have, as a result, seen something of a burst of new brocket species being described. It is quite probable that a similar taxonomic explosion may be in line for the genus Odocoileus, with both of the currently recognised species including a number of subspecies and each being of suspect monophyly. Matters are further complicated by the possibility of hybridisation between the two 'species'.


Gilbert, C., A. Ropiquet & A. Hassanin. 2006. Mitochondrial and nuclear phylogenies of Cervidae (Mammalia, Ruminantia): systematics, morphology, and biogeography. Molecular Phylogenetics and Evolution 40 (1): 101–117.

Pitra, C., J. Fickel, E. Meijaard & P. C. Groves. 2004. Evolution and phylogeny of Old World deer. Molecular Phylogenetics and Evolution 33: 880–895.

Conus jaspideus or Conasprella jaspidea, Take Your Pick

Live Conasprella jaspidea, copyright Anne DuPont.

Cone shells are one of the classic varieties of tropical sea shells, perhaps only rivalled in their familiarity with the general public by cowries and conches. Over 800 species of the family Conidae have been described from around the world. The specimen above represents one of these species, going by the name of Conasprella jaspidea or Conus jaspideus. The alternatives reflect the conflict between those who would treat all cone shells as belonging to a single genus Conus, or those who would divide them between multiple genera (Conasprella jaspidea is the name used for this species by Puillandre et al., 2014). One 2009 classification went so far as to divide the cone shells between 89 genera in five separate families, which does seem perhaps a little excessive. Among other features, Conasprella species differ from Conus sensu stricto in having a higher spire to the shell.

The type specimen of Conasprella jaspidea, copyright MHNG.

Conasprella jaspidea is found in coastal sections of the western Atlantic between Florida and the area of Rio de Janeiro. It is a medium-sized shell, reaching about three centimetres in length. Whorls of the spire are marked by distinct shoulders, and the body whorl is ornamented by spiral cords. The colour of the shell is white, orange or brown with darker brownish or violet spots. Shells of C. jaspidea may vary in texture from granular to smooth. These variants were initially recognised as distinct species or subspecies Conus jaspideus and C. verrucosus but, not only can both forms be found intermixed within a single population, the difference between them may be simply a question of the degree of wear a shell has been exposed to (Santos Gomes 2011).

Like other cone shells, Conasprella jaspidea is venomous with the radula bearing a single functional tooth modified into a short of hypodermic needle for injecting venom. Species of Conasprella are vermivorous (that is, they feed on worms). Feeding by a live individual of C. jaspidea was observed in an aquarium by Santos Gomes (2011). Photographs therein show the individual ingesting a polychaete worm that was perhaps not too much shorter in length than the cone shell itself; the process of feeding (from the initial strike with the radula to completion of ingestion) took about eighteen minutes from start to finish.


Puillandre, N., T. F. Duda, C. Meyer, B.M. Olivera & P. Bouchet. 2014. One, four or 100 genera? A new classification of the cone snails. Journal of Molluscan Studies 81: 1–23.

Santos Gomes, R. dos. 2011. Conus jaspideus (Mollusca: Neogastropoda: Conoidea) on the Brazilian coast. Journal of the Marine Biological Association of the United Kingdom 91 (2): 531–538.

Darklings, Tok Toks and Pie-dishes

False wireworm beetle Gonocephalum sp., copyright EBKauai.

It has been noted to the point of cliché that the Creator has an inordinate fondness of beetles. Even within the massive range of beetle diversity, though, certain families stand out as particularly diverse. One such family is the Tenebrionidae, with over twenty thousand known species worldwide. The family is sometimes referred to as the darkling beetles but no one vernacular name is really sufficient for this group. Not only are tenebrionids taxonomically diverse, they are morphologically diverse, varying from long-legged and elongate to hemi-spherical and robust, from smooth and shining to ornate and hairy, from dull-coloured and retiring to bright and striking. Habits vary from detritivorous to xylophagous (feeding on decaying wood) to herbivorous to mycetophagous, with even a few predators. Larvae of some species are of economic significance as pests: the false wireworms feed on the roots of crops or lawns, while mealworms and flour beetles attack stored products (mealworms are, of course, also used as pet food and occasionally even as human food). Several species live as inquilines of social insects such as ants or termites. The highest diversity of tenebrionids is in relatively arid regions; some species, such as the tok tok beetles of southern Africa and the pie-dish beetles of Australia, are familiar sights in such habitats.

Pie-dish beetle Helea sp., copyright Australian Museum.

With such high diversity, it is not easy to define this group without encountering exceptions, but generally tenebrionids have the antennae eleven-segmented and inserted below lateral expansions of the genae. The procoxal cavities are usually closed externally, and the legs of most species have a 5-5-4 tarsal formula. The first three sternites of the abdomen are fused (Kergoat et al. 2014). Several subfamilies are recognised, but they are commonly grouped into three clusters known as the lagrioid, pimelioid and tenebrionoid branches of the family (Matthews & Bouchard 2008). Many members of the lagrioid and tenebrionoid branches possess well-developed defensive glands in the abdomen. The rear sternites of the abdomen in these species are hinged on the sides rather than along the midline as in more primitive forms, allowing the abdomen to expand as the gland reservoirs fill with a repugnant fluid that can be expelled when required. Many larger tenebrionids have a tendency to walk with their rear ends tilted upwards, ready to unleash at a moment's notice.

Allecula rhenana, copyright Stanislav Krejčík.

Members of the pimelioid branch, including the subfamilies Pimeliinae and (possibly) Zolodininae, lack abdominal defensive glands. In many parts of the world, pimelioids are the dominant tenebrionids in dry habitats. The lagrioid branch includes the single subfamily Lagriinae, defined by features of the genitalia. Matthews & Bouchard (2008) also listed the small subfamily Phrenapatinae in this branch but a molecular phylogenetic analysis of the family by Kergoat et al. (2014) placed this latter subfamily in the tenebrionoid branch. The tenebrionoid branch also includes the Tenebrioninae, Diaperinae, Alleculinae and Stenochiinae, though monophyly of the Tenebrioninae and Diaperinae is uncertain (Kergoat et al. 2014). Diaperines include a number of shiny, sometimes strikingly coloured species; members of the tribe Leiochrinini look more like ladybeetles of the Coccinellidae than typical tenebrionids. The Tenebrioninae include such notable members as the false wireworms of the tribe Opatrini, the mealworms of the Tenebrionini and the flour beetles of the Triboliini. Finally, the Alleculinae are a distinctive group of often relatively soft-bodied tenebrionids readily distinguished from other members of the family by their pectinate claws; in some older classifications, alleculines were treated as a separate family of their own.


Kergoat, G. J., L. Soldati, A.-L. Clamens, H. Jourdan, R. Jabbour-Zahab, G. Genson, P. Bouchard & F. L. Condamine. 2014. Higher level molecular phylogeny of darkling beetles (Coleoptera: Tenebrionidae). Systematic Entomology 39: 486–499.

Matthews, E. G., & P. Bouchard. 2008. Tenebrionid Beetles of Australia. Australian Biological Resources Study.


The world is home to a wide variety of leafhoppers, both in terms of number of species and range of morphological disparity. One of the more diverse leafhopper families is the Delphacidae, including over two thousand species from around the globe. Delphacids are relatively small leafhoppers that are easily distinguished from other families by the possession of a large movable spur at the end of the tibia of the hind leg. I can't say as I know what the function of this spur is, but similar structures in other insect groups may be used for grooming.

Brown leafhoppers Nilaparvata lugens, from ICAR. The individual on the right is a long-winged disperser, the one on the left is a flightless brachypter.

Delphacids feed on the phloem of their host plants; the greater number of species are associated with monocots such as grasses. A number of species are significant economic pests; perhaps the most infamous are the brown leafhopper Nilaparvata lugens and white-backed leafhopper Sogatella furcifera which attack rice. They feed at the base of rice plants, causing the formation of round, yellow patches that soon dry up and turn brown, a condition known as 'hopper burn'. Death of the entire plant will often follow. As well as the direct damage from feeding, these leafhopper species also transmit viruses that further impact yields. Historically, numerous famines have been blamed on leafhopper outbreaks, such as the Kyoho famine of 1732 that saw rice production reduced to only 10% of its previous level. Estimates of the number of people affected by the famine seem to vary widely—according to Wikipedia, the official death toll was a bit more than twelve thousand people, but estimates of the actual number of fatalities range well in excess of 150,000. In more recent years, leafhopper outbreaks may be exacerbated by indiscriminate fertiliser and pesticide use, with the latter reducing competition for the hoppers from other insects or predators.

Delphacids (and many other leafhoppers) commonly exhibit polymorphism in wing development with both flying macropterous and flightless brachypterous forms occuring in a single population. The question of macroptery vs brachyptery is an environmental one. If a developing delphacid receives sufficient nitrogen then it will develop into a flightless adult, remaining in the place of its birth to continue to benefit from the good feeding conditions there. But if feeding conditions become degraded and the developing nymph is deprived of nitrogen then it will develop into a fully-winged adult that can leave its home in search of more favourable conditions elsewhere. Because of their small size, migrating delphacids may be carried long distances by the winds. In the case of pest species, this phenomenon of migration further exacerbates the problem of control as hopper populations from different countries are regularly mixed, increasing genetic diversity and resistance to varying control methods.


Urban, J. M., C. R. Bartlett & J. R. Cryan. 2010. Evolution of Delphacidae (Hemiptera: Fulgoroidea): combined-evidence phylogenetics reveals importance of grass host shifts. Systematic Entomology 35: 678–691.

When the Wolf Breaks Wind

Common puffballs Lycoperdon perlatum, copyright H. Krisp.

In an earlier post, I described the way in which the 'gasteromycetes' of historical fungal classifications have come to be expunged as a category. The enclosure of spore-producing structures within a contained fruiting body such as a puffball, instead of exposed on a membrane such as on the underside of a mushroom cap, is something that has evolved many times in fungal history. One possible suggestion for why this may occur is as a protection against moisture loss, allowing the fungus to thrive in drier or more exposed habitats than before.

In that earlier post, I also mentioned off-hand that one of the best known groups of 'gasteromycetes', the puffballs of the Lycoperdaceae, are in fact close relatives of some of the best known typical mushrooms in the Agaricaceae. Indeed, it appears that recent authors may go so far as to synonymise the two families. Puffballs emerge as globular fruiting bodies that become packed with spores as they mature, until one or more openings develop in the external skin of the fruiting body and allow the pores to escape. Supposedly many puffballs are quite edible if collected before the spores begin to develop, though I've never tried myself. Particularly sought in this regard is the giant puffball Calvatia gigantea whose fruiting bodies grow particularly large; supposedly, examples have been found over a metre in diameter and weighing up to twenty kilogrammes.

Giant puffball Calvatia gigantea, copyright Alan Wolf.

Dispersal of spores from puffballs may be achieved in a number of ways. In species found in habitats with more regular rainfall, such as species of the genus Lycoperdon, spores are spread by 'boleohydrochory' (Gube & Dörfelt 2011). '-Chory' means dispersal, '-hydro-' obviously means water, 'boleo-' I think may mean something like 'throw'. The puffball opens through a hole in the top, and drops of rain (or other sources of pressure such as being tapped by an animal) cause a puff of spores to be squeezed out. The water may then carry the spores away. The name Lycoperdon, as it happens, literally translates as 'wolf fart', and this is another one of those names I am completely at a loss to explain. The 'fart', obviously, refers to the appearance of the spore puffs, but what on earth do they have to do with wolves?

Tumbling puffballs Bovista pila, copyright Dan Molter.

Other puffballs may spread their spores via 'anemochory', dispersal by wind. This is particularly the case with species found in drier habitats. Some species, such as some members of the genus Bovista, exhibit a variation on this called 'geanemochory' in which the entire puffball becomes detached and blown about by the wind, with the spores escaping through openings in the external shell like pepper being shaken from a pepper-pot. Differences in dispersal method between puffball species are generally reflected by differences in their spore morphology. Hydrochorous species usually have strong ornamentation, with the outside of the spore being covered with warts or the like. These warts provide more surface area for the water to catch onto; they may also help prevent the spores from clumping together. In contrast, anemochorous species have spores that are smooth, making them more streamlined for being blown through the air or, particularly in the case of geanemochorous species, making them less likely to become trapped by hyphae or other structures inside the fruiting body itself and so facilitating their escape.


Gube, M., & H. Dörfelt. 2011. Gasteromycetation in Agaricaceae s. l. (Basidiomycota): morphological and ecological implementations. Feddes Repertorium 122 (5–6): 367–390.

The Forams that Bind

Cross-section of Fabiania cassis, from BouDagher-Fadel (2008).

Here we see an example of Fabiania. Fabiania is a genus of foraminiferan known from the Eocene epoch that could reach a relatively large size as forams go, up to several millimetres across (nowhere near as large as some that I've covered on this site, maybe, but still respectable). It had a conical test with a rounded apex and a deeply excavated centre; depending on growing conditions, individual Fabiania might be a regular or a flattened cone. In its early stage, Fabiania had two globose thick-walled and perforate chambers; later chambers were cyclical and divided by horizontal and vertical partitions. The aperture of the test was a single row of pores opening into the large umbilicus. The wall of the test was thick and calcareous, and covered with coarse perforations on the upper side of the cone (BouDagher-Fadel 2008; Loeblich & Tappan 1964).

Fabiania lived in association with coral reefs, often preferring the undersides of corals and other sheltered locations. It was primarily found around the mid-depths, not too close to the water's surface but also not too deep (Bosellini & Papazzoni 2003). I've referred in an earlier post to another group of coral-encrusting forams, the acervulinids. Because reef forams tend to be cryptic (in more exposed parts of the reef they tend to get out-competed by coralline algae), and are often variable in morphology making them taxonomically difficult, they tend to be less studied than the reef's more prominent components. However, forams may play a not so insignificant role in developing the reef's structure, helping to bind the reef in place.


Bosellini, F. R., & C. A. Papazzoni. 2003. Palaeoecological significance of coral-encrusting foraminiferan associations: a case-study from the Upper Eocene of northern Italy. Acta Palaeontologica Polonica 48 (2): 279–292.

BouDagher-Fadel, M. K. 2008. The Cenozoic larger benthic foraminifera: the Palaeogene. Developments in Palaeontology and Stratigraphy 21: 297–418.

Loeblich, A. R., Jr & H. Tappan. 1964. Treatise on Invertebrate Paleontology pt C. Protista 2. Sarcodina, chiefly "thecamoebians" and Foraminiferida vol. 1. The Geological Society of America and The University of Kansas Press.