Field of Science

Dealing with a Clingy Male

Diving beetles of the family Dytiscidae are a distinctive component of the freshwater environment in most regions of the world. They have an oval, streamlined body form and powerful hind legs, usually with fringes of stiff setae, that are ill-suited for movement on land but make them adept swimmers. They are also almost always capable fliers, allowing them to find their way to water bodies of any size from large lakes to small, temporary pools. Both adults and larvae are active hunters, preying on other aquatic arthropods or even small vertebrates. Most diving beetles are fairly dull in coloration but exceptions are found among members of the tribe Aciliini.

Sunburst diving beetle Thermonectus marmoratus, from Insectarium de Montréal, René Limoges.


Members of the Aciliini are moderately sized diving beetles, generally between one or two centimetres in length. Dorsally they have a yellow to red base coloration with contrasting dark markings. The hind legs are robust with the hind tibia short and broad. Males have the base of the tarsus of the front legs broadened into a round palette with setae on the underside modified into sucking discs, used to hang onto the females when mating; this discs may be present on the tarsus of the mid pair of legs as well. They are strong swimmers, often venture=ing into the open waters of lakes and pools, and contrast with other diving beetles in that they may be found in pools lacking submerged vegetation (Roughley & Larson 2001; Bergsten & Miller 2006). Larvae have a distinctive arched body shape with a small head (Bukontaite et al. 2014), kind of shrimp-like, and also tend to be more pelagic than the larvae of other diving beetles. Females have gonocoxae (the appendages at the end of the abdomen that function as the ovipositor) that are relatively long with a broadened, spoon-like ending (Miller 2001); these are used to insert eggs into damp moss or under loose bark of vegetation lying just above the waterline. There is usually just one generation per year and adults in cold regions overwinter in larger water bodies that remain unfrozen.

Alternate morphs of female Graphoderus zonatus with granular (left) and smooth elytra, from Holmgren et al. (2016).


Perhaps the most intriguing aspect of aciliin diving beetles regards their sexual dimorphism. As noted above, males have a set of suckers on the fore legs for hanging onto females when mating. However, females of some species have sculpted elytra rather than the smooth elytra of males, such as a granular surface in Graphoderus species or long, setose sulci in female Acilius. The uneven surface produced by these features presumably functions to reduce the efficacy of the males' suckers, allowing the females more control when selecting a mate. That such a conflict exists is supported by the observation that the more developed the males' sucker arrays in a population, the more likely the females are to have repellent sculpturing. Males of some diving beetle species have been observed grabbing at any female they encounter, followed by the female swimming rapidly and erratically in an attempt to shake the male off or knock him off against the substrate or objects in the water (Miller 2003). Where this becomes really interesting is that some species have dimorphic females with some females in the population having sculpted elytra whereas others are smooth. What could be the reason for such variation? The presence of both forms in the population suggests that neither has a complete advantage over the other. It may be that smooth-backed females trade reduced defenses for improved swimming ability. Alternatively, a defensive female may be able to ensure that only the strongest and most resilient males can mate with her, but runs the risk of not mating at all if she never encounters a male who can overcome her defenses. A less defensive female may be more vulnerable to any male she encounters but at least she's bound to be fertilised at some point.

REFERENCES

Bergsten, J., & K. B. Miller. 2006. Taxonomic revision of the Holarctic diving beetle genus Acilius Leach (Coleoptera: Dytiscidae). Systematic Entomology 31: 145–197.

Bukontaite, R., K. B. Miller & J. Bergsten. 2014. The utility of CAD in recovering Gondwanan vicariance events and the evolutionary history of Aciliini (Coleoptera: Dytiscidae). BMC Evolutionary Biology 14: 5.

Holmgren, S., R. Angus, F. Jia, Z. Chen & J. Bergsten. 2016. Resolving the taxonomic conundrum in Graphoderus of the east Palearctic with a key to all species (Coleoptera, Dytiscidae). ZooKeys 574: 113–142.

Miller, K. B. 2003. The phylogeny of diving beetles (Coleoptera: Dytiscidae) and the evolution of sexual conflict. Biological Journal of the Linnean Society 79: 359–388.

Roughley, R. E., & D. J. Larson. 2001. Dytiscidae Leach, 1815. In: Arnett, R. H., Jr & M. C. Thomas (eds) American Beetles vol. 1. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia pp. 156–186. CRC Press: Boca Raton.

Atropacarus

The little guy pictured above (photo copyright Scott Justis) is a representative of the box mite genus Atropacarus, members of which can be found in most parts of the world. Atropacarus is a genus of the Phthiracaroidea, a group of box mites characterised by the plates on the underside of body being relatively wide, in contrast to the narrow ventral plates of its sister group, the Euphthiracaroidea (members of which have featured on this site before: here and here). The difference in configuration of these plates reflects a difference in the way that the body is contracted to allow legs and prosoma to be withdrawn beneath the protective cover of the notogaster. In euphthiracaroids, the sides of the notogaster are contracted inwards; in phthiracaroids, the ventral plates of the body are lifted upwards (Schmelzle et al. 2015).

The classification of phthiracaroids is subject to conflict with two main systems in the recent literature. In one, championed by the Polish acarologist Wojciech Niedbała, the phthiracaroids are divided between two families with Atropacarus in the Steganacaridae. Species of Atropacarus have the surface of the notogaster extensively covered with dimples. The dorsal seta on the tibia of the fourth leg is short and closely associated with a solenidion (a type of specialised sensory hair). The setae of the genital plate are arranged in a more or less straight row along the inner margin of the plate with the fifth and sixth setae further apart than the fourth and fifth (Niedbała 1986). Niedbała divides Atropacarus between two subgenera. In Atropacarus sensu stricto, there are sixteen or more pairs of setae on the notogaster and the second adanal seta is moved inwards on the ano-adanal plate to form a more or less straight line with the anal setae. In Hoplophorella, there are fifteen pairs of setae on the notogaster and the second adanal seta is distinctly laterally placed relative to the anal setae.

The super-hairy Atropacarus niedbalai, from Liu & Zhang (2013). Scale bar = 100 µm.


In the competing system, used for instance by Subías (2019), Atropacarus and Hoplophorella are treated as distinct genera and each is in turn divided into subgenera by the number of setae on the ano-adanal plate. To a certain extent, of course, the question of whether to treat Atropacarus and Hoplophorella as genera or subgenera is arbitrary. Nevertheless, this arguably cosmetic distinction does relate to an underlying difference in theory. The classification of phthiracaroids used by Subías (2019) is a largely diagnostic one, inspired by a desire to facilitate specimen identifications. Niedbała's classification, in contrast, is intended to reflect phylogenetic relationships. Simple setal counts may be convenient when composing keys but one might question its overall phylogenetic significance. Neotrichy (increases in setal count by multiplication of the original setae) is not uncommon in phthiracaroids, particularly on the notogaster. Setal counts may vary between individuals of a single species and overall neotrichy reaches an extreme in the New Zealand species Atropacarus niedbalai. In this species, the basic count of fifteen or sixteen pairs of notogastral setae has been increased to 109 or 115 pairs, with further neotrichy on the prodorsum and ventral plates (Liu & Zhang 2013). Subías (2019) defends his choice of classification by arguing that Niedbała's key features are often difficult to discern. I sympathise with the difficulty but, as a wise man once said, species are under no obligation to evolve with regard to the convenience of taxonomists.

REFERENCES

Liu, D., & Z.-Q. Zhang. 2013. Atropacarus (Atropacarus) niedbalai sp. nov., an extreme case of neotrichy in oribatid mites (Acari: Oribatida: Phthiracaridae). International Journal of Acarology 39 (6): 507–512.

Niedbała, W. 1986. Système des Phthiracaroidea (Oribatida, Euptyctima). Acarologia 27 (1): 61–84.

Schmelzle, S., R. A. Norton & M. Heethoff. 2015. Mechanics of the ptychoid defense mechanism in Ptyctima (Acari, Oribatida): one problem, two solutions. Zoologischer Anzeiger 2015: 27–40.

Subías, L. S. 2019. Nuevas adiciones al listado mundial de ácaros oribátidos (Acari, Oribatida) (14a actualización). Revista Ibérica de Aracnología 34: 76–80.

The Font of the Placentals

The large-scale incorporation of molecular data into phylogenetics over the last few decades has caused a revolution in our understanding of life's evolution. Taxa whose interrelationships were previously regarded as intractable have been opened up to study, and many of our previous views on relationships have been forced to shift. Because conflict always makes for a good story, certain cases of the latter have become causes celebres, receiving extensive attention in both the technical and popular literature. One of these subjects of particular interest, not surprisingly, involves the relationships of the living orders of mammals.

Reconstruction of Arctostylops steini by Brian Regal, from Janis et al. (1998). The arctostylopids are a Palaeocene to Eocene group of mammals of uncertain affinities but probably belonging somewhere in the Boreoeutheria.


A lot of this attention has focused around the revelation of the Afrotheria, a grouping of animals (tenrecs, elephant shrews, hyraxes, aardvarks, elephants and manatees) with likely African origins that was completely unsuspected by studies based on morphological data only but which molecular studies have identified with ever-increasing levels of support. Recent molecular studies of placental phylogeny have agreed on three basal divisions within the placental mammals: the Afrotheria, the Xenarthra (armadillos, anteaters and sloths, a grouping that was recognised even before the advent of molecular data), and the remaining placentals in the largest of the three, the Boreoeutheria.

To the best of my knowledge, the Boreoeutheria is a clade that has also so far been supported by molecular data only with no morphological features yet recognised as defining the group. Nevertheless, its support can be considered as well established. The name Boreoeutheria refers to the clade's likely northern origins in contrast to the more southern distribution of the other two. Within the Boreoeutheria, molecular studies indicate a basal divide between the Euarchontaglires on one side and the Laurasiatheria on the other. The Euarchontaglires include the primates and rodents (as well as a handful of smaller orders). The Laurasiatheria include the Eulipotyphla, a group of insectivorous mammals including shrews, moles and hedgehogs, as sister to a clade containing bats, carnivorans, perissodactyls and artiodactyls.

Molecular phylogeny of mammals, from Springer et al. (2004) (note that not all branches shown in this tree are supported by all studies).


This all has interesting ramifications for the early evolution of placentals. There is an extensive fossil record of mammals from the Palaeocene, the epoch of time immediately following the end of the Cretaceous. However, most of these mammals do not belong to the orders alive today and their exact relationships to living mammals remain open to debate. The molecule-induced shake-up of pacental relationships just increased this uncertainty: for instance, the interpretation of a given group of fossil mammals as close to the common ancestry of perissodactyls and elephants rather goes out the window when perissodactyls and elephants are no longer thought to be closely related. And detailed studies that may resolve these issues remain few and far between. One of the most notable analyses in recent years has been that by Halliday et al. (2017) which covered most of the well-preserved placentals and their close relatives from the Cretaceous and Palaeocene periods. However, it is difficult to say just what to make of their results. The unconstrained analysis of their data presents results that remain deeply inconsistent with the molecular tree. Conversely, constraining the analysis to more closely match the molecular data provides results that are intriguing but difficult to accept at face value; I suspect they may be artefacts of the algorithm forcing taxa into the least unacceptable position for inadequate data. Suggesting that pangolins are the last specialised survivors of a broad clade of condylarths, pantodonts, notoungulates and creodonts is... I suppose not a priori impossible, but definitely a big call. A later analysis based on an expanded version of the same data set by Halliday et al. (2019) irons out some of the kinks but still fails to resolve the base of the Boreoeutheria beyond a massive polytomy of 25 branches (an icosipentatomy?). The Euarchontaglires are recovered as a clade but not the Laurasiatheria or any of its molecular subgroups above the ordinal level. And while some of the newer analysis' placements may seem like an improvement (notoungulates are placed as the sister to litopterns instead of hanging out with pangolins), others may still raise an eyebrow (mesonychids are associated with carnivorans but viverravids and miacids are not).

As always, the best answer to this conundrum is likely to involve more research. While researching this post, I did come across comments from people suggesting issues with the Halliday et al. data. Frankly, for a data set of this size (involving 248 taxa and 748 characters in the 2019 paper), it would be incredible were it otherwise. I know from my own experience that as you add more characters and taxa to a phylogenetic analyses, the challenge of keeping everything in line rises exponentially, and the data sets I've dealt with have been nowhere near the size of this one. Nevertheless, it's a start. And we can but hope that even those who find fault with it ultimately take it as inspiration to themselves do better.

REFERENCES

Halliday, T. J. D., M. dos Reis, A. U. Tamuri, H. Ferguson-Gow, Z. Yang & A. Goswami. 2019. Rapid morphological evolution in placental mammals post-dates the origin of the crown group. Proceedings of the Royal Society of London Series B—Biological Sciences 286: 20182418.

Halliday, T. J. D., P. Upchurch & A. Goswami. 2017. Resolving the relationships of Paleocene placental mammals. Biological Reviews 92 (1): 521–550.

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 2nd 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.