Williamsita

A while back, I wrote a post about the crabronid wasp genus Podagritus. This time, I'm going to cover another crabronid genus found here in Australia: Williamsita.

Williamsita sp., copyright David Francis.

Like Podagritus, Williamsita species are boldly coloured wasps, typically mostly black with contrasting yellow or orange markings. They differ from Podagritus species in being more robust with the base of the gaster not notably pedunculate. Other distinguishing features include the presence of distinct foveae (pits) against the margins of the eyes (occasionally less distinct in males), thirteen-segmented antennae in males, and a pygidial plate in both sexes that is narrowed and concave in females, quadrate in males. Williamsita species also do not have the palps reduced as in Podagritus, instead having the more typical arrangement of six segments in the maxillary palps and four segments in the labial palps (Bohart & Menke 1976).

To date, eleven species have been recognised in the genus Williamsita (Leclercq 2006). Most are found in Australia with a single species each known from New Caledonia and Vanuatu. Leclercq (1950) suggested dividing the genus between two subgenera with all species except the New Caledonian type species W. novocaledonica forming a subgenus Androcrabro. Features supporting the latter taxon included the presence of ventral notches on one or more segments of the antennae in males. However, Leclercq later suggested abandoning such a formal division, questioning its significance (Leclercq 2006). The Australian species of Williamsita are, nevertheless, distinct from the two insular species in being marked with much stronger punctation over the body.

Most Williamsita species remain little seen and poorly known. However, breeding habits have been recorded for two Australian species, W. bivittata and W. tasmanica (Maynard & Fearn 2021; McCorquodale et al. 1989). Both these species nest in branching holes in rotting wood, either commandeering burrows left by wood-boring insects or excavating their own. Prey consists of larger flies such as blow flies or soldier flies which were carried back to the nest by the wasp running with the fly carried below the body. Up to six paralysed flies might be placed lying on their backs in a nest cell with an egg laid across the 'throat' (i.e. at the joint between head and thorax) of one of the flies. The cell would then be closed with a plug of woody frass. McCorquodale et al. (1989) recorded W. bivittata constructing several such cells in a series along a single tunnel, whereas Maynard & Fearn (2021) found W. tasmanica more likely to place a single cell in a side-branch. As both observations were limited to a single location in a single season, though, one might reasonably question whether these represent true differences in species behaviour or were determined by available conditions. There's a limit to how deep a Williamsita can burrow.

REFERENCES

Bohart, R. M., & A. S. Menke. 1976. Sphecid Wasps of the World. University of California Press: Berkeley.

Leclercq, J. 1950. Sur les crabroniens orientaux et australiens rangés par R. E. Turner (1912–1915) dans le genre Crabro (subgenus Solenius). Bulletin et Annales de la Société Entomologique de Belgique 86 (7–8): 191–198.

Leclercq, J. 2006. Hyménoptères crabroniens d'Australie du genre Williamsita Pate, 1947 (Hymenoptera: Crabronidae). Notes Fauniques de Gembloux 59 (2): 115–119.

Maynard, D., & S. Fearn. 2021. Ecological and behavioural observations of a nesting aggregation of the endemic Tasmanian digger wasp Williamsita tasmanica (Smith, 1856) (Hymenoptera: Crabronidae: Crabroninae). Papers and Proceedings of the Royal Society of Tasmania 155 (1): 43–50.

McCorquodale, D. B., C. E. Thomson & V. Elder. 1989. Nest and prey of Williamsita bivittata (Turner) (Hymenoptera: Sphecidae: Crabroninae). Australian Entomological Magazine 16 (1): 5–8.

Opening Dors

My current dayjob mostly revolves around identifying and counting dung beetles. When Europeans settled Australia, they brought their farm animals with them. Unfortunately, the large piles of dung produced by cattle and horses proved rather daunting to native scavengers used to the more compact droppings of kangaroos and possums. And if you've ever experienced an Australian summer, you'll know that flies are definitely a thing. To help with this situation, Australia has had a long-running programme introducing exotic dung beetles that are better able to clean up after livestock. Most of these are members of the typical dung beetle family Scarabaeidae but one species, Geotrupes spiniger, represents a different subgroup of the superfamily Scarabaeoidea. These are the earth-boring dung beetles or dor beetles of the Geotrupidae.

Dor beetle Geotrupes spiniger, copyright Udo Schmidt.
The geotrupids are medium-sized to very large beetles, ranging in size from half a centimetre to 4.5 cm in length (Jameson 2002). Like many other members of the Scarabaeoidea, they have broad fore legs used for digging. Their short, eleven-segmented antennae end in the asymmetrical club typical of scarabaeoids but they may be distinguished from other families in that the basal segment of the three-segmented club is expanded to form a 'cup' against which the other segments may be tightly closed. The body of geotrupids is strongly convex, and is smooth and shiny dorsally but hairy underneath. In many species, the males may bear elaborate horns and/or processes on the head and pronotum.

Male Taurocerastes patagonicus, copyright Nicolás Lavandero.

Despite their size, geotrupids are secretive animals, spending most of their time in burrows underground (which may be up to three metres in depth) and usually only emerging at night. Various species feed on animal dung or decaying matter; some feed on subterranean fungi. In at least some species, eggs are laid in brood chambers within the parent's home burrow and multiple life stages may share a single burrow. Burrows may also be shared between multiple adults when conditions demand. Though adults do not directly tend to larvae, they may stock brood chambers with food supplies. In some Australian species of the subfamily Bolboceratinae, females lay a single gigantic egg at a time that may be up to 56% the size of its layer (Houston 2011). Larvae hatching from such an egg are able to develop right through to maturity without feeding.

Adult geotrupids produce a stridulating noise when disturbed which is the origin of the alternate vernacular name of "dor beetle" ("dor" being an old word for a buzzing insect). Larvae may or may not be capable of stridulation, depending on the species.

Male Blackburnium rhinoceros, copyright Edward Bell.

The classification of geotrupids is the subject of ongoing investigation. A recent classification divides the family between three subfamilies, the widespread Geotrupinae and Bolboceratinae and the South American Taurocerastinae. Morphological differences between these subfamilies, particularly at the larval stage, have lead some researchers to question whether the Geotrupidae in the broad sense represents a monophyletic group. Molecular analyses thus far seem ambiguous; an analysis by McKenna et al. (2015) placed geotrupids as part of a polytomy near the base of the scarabaeoids. As an aside, my supervisor recently asked myself and a retired colleague whether Geotrupes spiniger was the only species of geotrupid found in Australia. I replied "yes", our colleague responded "no". Our conflict, of course, was based on whether Australia's wide diversity of Bolboceratinae contributed to the count.

REFERENCES

Houston, T. F. 2011. Egg gigantism in some Australian earth-borer beetles (Coleoptera: Geotrupidae: Bolboceratinae) and its apparent association with reduction or elimination of larval feeding. Australian Journal of Entomology 50: 164–173.

Jameson, M. L. 2002. Geotrupidae Latreille 1802. In: Arnett, R. H., Jr, M. C. Thomas, P. E. Skelley & J. H. Frank (eds) American Beetles vol. 2. Polyphaga: Scarabaeoidea through Curculionoidea pp. 23–27. CRC Press.

McKenna, D. D., B. D. Farrell, M. S. Caterino, C. W. Farnum, D. C. Hawks, D. R. Maddison, A. E. Seago, A. E. Z. Short, A. F. Newton & M. K. Thayer. 2015. Phylogeny and evolution of Staphyliniformia and Scarabaeiformia: forest litter as a stepping stone for diversification of nonphytophagous beetles. Systematic Entomology 40: 35–60.

Platybunus: the Wide-Eyed Harvestmen of Europe

The western Palaearctic region (that is, Europe and the immediately adjacent parts of Asia and northern Africa) is home to a diverse and distinctive fauna of harvestmen. Among the various genera unique to this part of the world are the forest- and mountain-dwellers of the genus Platybunus.

Platybunus pinetorum, copyright Donald Hobern.

Platybunus species are moderate-sized long-legged harvestmen of the family Phalangiidae, the central body in larger individuals being about eight millimetres long (Martens 1978). Their most characteristic feature is a relatively large eye-mound, distinctly wider than long and occupying a large section of the anterior carapace. As with other European phalangiids, they eye-mound is ornamented with a row of denticles each side though the body lacks denticles over the remainder of the dorsum. The body is often comparatively slender, tapering towards the rear (particularly in males), and is marked on the dorsum by a darker median band. The pedipalps have a pair of well-developed setose apophyses on the inner distal ends of the patella and tibia, and a series of long spine-like tubercles on the underside of the femur. These tubercles presumably function in the capture of prey, forming a basket that can be closed around the harvestman's victims. External sexual dimorphism in Platybunus is fairly minimal though females are overall larger and fatter. The penis is notably long and slender with a relatively small glans, offset from the shaft by a more or less marked constriction.

Platybunus bucephalus, copyright Adrian Tync.

Martens (1978) recognises four species of Platybunus found in higher altitude regions of central Europe with the species P. bucephalus and P. pinetorum occupying much of the genus' range. Platybunus bucephalus may be distinguished from P. pinetorum by, among other features, its relatively shorter legs. Platybunus pallidus is endemic to the Carpathians, and the tiny P. alpinorelictus inhabits the Garda Mountains of northern Italy. Another species, P. anatolicus, was described from Turkey by Roewer (1956)*. In general, Platybunus species inhabit alpine and subalpine forests, being found among the herbaceous undergrowth, under bark or on rock faces. Where their ranges overlap, P. bucephalus is more accustomed to extending beyond the forest margins than P. pinetorum and may be found above the tree-line. In recent years, the range of P. pinetorum has extended northwards, being first recorded from the UK in 2010 and Sweden in 2015 (Fritzén et al. 2015). At least some populations of P. pinetorum are capable of reproducing parthenogenetically and this may have played a part in its spread.

*Platybunus mirus was described by Loman (1892) on the basis of two male specimens that supposedly came from Sumatra. Though the identity of this species has never been resolved (Loman's illustration of the penis is at least suggestive of a true Platybunus), the claimed locality seems almost certain to be an error of some kind.

The internal classification of the Phalangiidae remains in need of further investigation. Platybunus has been recognised by some authors as forming a subfamily Platybuninae with a cluster of other western Palaearctic genera bearing similar ventrally spined pedipalps (Zhang & Zhang 2012). However, other authors have not separated this group from the subfamily Phalangiinae. The platybunines may represent a phylogenetically coherent grouping, or their shared features may reflect adaptations to a similar life style. The genital morphology of Platybunus is recognisably distinct from that of other platybunines which may argue against any relationship (Martens 1978). On the other hand, platybunines might possibly be distinguished from phalangiines by the chemical composition of their repugnatorial gland secretions (Raspotnig et al. 2015). A formal analysis of the family's evolution would be a welcome advance.

REFERENCES

Fritzén, N. R., V. Rinne, M. Sunhede, A. Uddström, S. Van de Poel & P. De Smedt. 2015. Platybunus pinetorum (Arachnida, Opiliones) new to Sweden. Memoranda Soc. Fauna Flora Fennica 91: 37–40.

Loman, J. C. C. 1892. Opilioniden von Sumatra, Java und Flores. In: M. Weber (ed.) Zoologische Ergebnisse einer Reise in Niederländisch Ost-Indien vol. 3 pp. 1–26, pl. 1. E. J. Brill: Leiden.

Martens, J. 1978. Spinnentiere, Arachnida: Weberknechte, Opiliones. Gustav Fischer Verlag: Jena.

Raspotnig, G., M. Schaider, P. Föttinger, V. Leutgeb & C. Komposch. 2015. Benzoquinones from scent glands of phalangiid harvestmen (Arachnida, Opiliones, Eupnoi): a lesson from Rilaena triangularis. Chemoecology 25: 63–72.

Roewer, C. F. 1956. Über Phalangiinae (Phalangiidae, Opiliones Palpatores). (Weitere Weberknechte XIX). Senckenbergiana Biologica 37 (3–4): 247–318.

Zhang, C., & F. Zhang. 2012. On the subfamilial assignment of Platybunoides (Opiliones: Eupnoi: Phalangiidae), with the description of a new species from China. Zootaxa 3190: 47–55.

Voley, Voley, Voley

Over a third of all living mammal species are rodents. In cooler regions of the Northern Hemisphere, the rodent fauna is often dominated by the Microtinae, the group of mouse-like rodents including voles and lemmings. And in North America, the most widespread of all microtine species is the eastern meadow vole Microtus pennsylvanicus.

Eastern meadow vole Microtus pennsylvanicus, copyright Gilles Gonthier.

The eastern meadow vole is found over most of Canada and a large part of the northern and eastern United States, with the subspecies M. p. chihuahuensis known from Chihuahua in northern Mexico. This species is about the size of a small rat, being from 14 to 20 cm in length with about three to six centimentres of that length being tail (Reich 1981). They are generally yellowish-brown in colour with black tips on the hairs though individuals vary significantly in brightness and shade. Western populations are supposed to be lighter in coloration than eastern, and southern individuals tend to be larger than northern. As an indication of this species' variability, Reich (1981) recognised 28 recognised subspecies.

Eastern meadow voles are primarily inhabitants of grasslands, with a preference for damper habitats, though they may also be found in woodlands. They mostly live in burrows underground, emerging to the surface to forage for food. Eastern meadow voles are generalist feeders, browsing on most available forms of low vegetation: grasses, sedges and herbs. When populations reach their peak, they may cause significant damage to woody plants by ringbarking their trunks. Individuals may seemingly be active at just about any time of day.

Eastern meadow vole in a state of danger, copyright David Allen.

Like other small rodents, meadow voles are short-lived animals with estimates of average lifespan ranging from just two or three months to ten to fourteen months (Reich 1981). Studies of movement patterns indicate that mature females generally maintain distinct, non-overlapping ranges whereas males range further and with less concern for others (Madison 1980). Mating behaviour appears generally promiscuous: males will range over the territories of multiple females and litters with mixed paternity are not uncommon (Boonstra et al. 1993). Paternal behaviour has been observed among eastern meadow voles in laboratory populations but all indications are that wild males do not remain with females after mating. Males often bear wounds indicative of intra-species conflict. These may be the result of males fighting over access to females but Madison (1980) suggested a potential alternative. Less dominant males might be more likely to attempt to approach females earlier or later in their oestrus cycle as the females are more likely to be guarded by dominant males when at their peak. While avoiding attacks from their dominant brethren, these minor males might find themselves violently rebuffed by a female who is just not yet in the mood.

After mating, gestation lasts for about three weeks, usually resulting in a litter of four to six babies. Weaning then takes place after about two weeks. Females forage far less while lactating than at other times. It might seem counter-intuitive for a female to reduce feeding when her energy demands are presumably at their peak but again Madison (1980) suggests an explanation: perhaps her energy needs are such that she simply lacks the capacity for extensive wandering. Young may potentially remain with their mother for some time after weaning but eventually they will be forced out of the parental burrow, leaving to face the wide world on their own. And when you're the size of a vole, that's a very wide world indeed.

REFERENCES

Boonstra, R., X. Xia & L. Pavone. 1993. Mating system of the meadow vole, Microtus pennsylvanicus. Behavioral Ecology 4: 83–89.

Madison, D. M. 1980. Space use and social structure in meadow voles, Microtus pennsylvanicus. Behavioral Ecology and Sociobiology 7: 65–71.

Reich, L. M. 1981. Microtus pennsylvanicus. Mammalian Species 159: 1–8.

Anchor Sponges

Sponges are, by their very nature, a challenging group taxonomically. At the macroscopic level, they are often amorphous and indeterminate in appearance. As one taxonomist complained in 1842 (as quoted in Hooper & Van Soest 2002): "there is so much that is in common to them, and each adapts itself so readily to circumstances and assumes a new mask, that it requires a tact, to be gained only by some experience, to recognize them under their guises; while we labour, perhaps in vain, to devise phrases which shall aptly portray to others the characteristics of objects that have no fixed shape, and whose distinctive peculiarities almost cheat the eye". Reliable identification typically requires the close examination of microscopic details, in particular the conformation and arrangement of the mineralised spicules that make up the skeleton of many sponges.

Myxilla incrustans, copyright B. E. Picton.

The Myxillidae are a family of marine sponges that, so far as we currently know, are most diverse in temperate and frigid waters. Like other members of the class Demospongiae, the most diverse of the recognised sponge classes, they have a skeleton of spicules constructed from silica. Different arrangements of spicules allow the body of the sponge to be divided into two layers. In the outer ectosoma, which can be thought of as the 'skin' of the sponge, elongate spicules are vertically radiating or placed in 'bouquet' arrangements with a palisade of vertical spicules surmounted by radiating clusters. These spicules generally have each end similar and may be smooth or spiky. In the inner choanosoma, within which are placed the feeding chambers of the sponge, elongate spicules are placed in a reticulate arrangement. These spicules generally have one end pointed and the other blunt.

Skeletal arrangements and individual spicules from various Myxillidae, from Hooper & Van Soest (2002).

Mixed in amongst these larger megasclere spicules are smaller microscleres that do not form part of the main structural skeleton, though presumably they do help hold the sponge body together. In myxillids, the microscleres generally take the form of anchorate chelae, small curved structures with incurved rounded prongs at each end. Members of the boreal genus Melonanchora have a mixture of chelae and a different type of microsclere shaped like a ribbed rugby ball (Santín et al. 2021). In the Indo-West Pacific genus Psammochela, growing sponges will also incorporate sand from the surrounding environment to supplement the microscleres (de Voogd 2012).

Growth habit of Myxillidae can vary from encrusting to massive to branching. The species Stelodoryx procera, found around the Azores, has a distinctive growth habit with a flattened main body at the end of an elongate stalk. On the whole, though individual species may be distinguished by growth habit, species within a single genus may differ greatly in form. For determining genera, examination of spicules is really the only way to go.

REFERENCES

Hooper, J. N. A., & R. W. M. Van Soest. 2002. Systema Porifera: A guide to the classification of sponges vol. 1. Kluwer Academic/Plenum Publishers.

Santín, A., M.-J. Uriz, J. Cristobo, J. R. Xavier & P. Ríos. 2021. Unique spicules may confound species differentiation: taxonomy and biogeography of Melonanchora Carter, 1874 and two new related genera (Myxillidae: Poecilosclerida) from the Okhotsk Sea. PeerJ 9: e12515.

Voogd, N. J. de. 2012. On sand-bearing myxillid sponges, with a description of Psammochela tutiae sp. nov. (Poecilosclerida, Myxillina) from the northern Moluccas, Indonesia. Zootaxa 3155: 21–28.

Succulent Orchids

With over 1200 known species found in Asia and Australasia, Dendrobium is one of the largest currently recognised genera of orchids. As with other examples of such 'super-genera', the question of how to best handle such a monster has been fiercely debated. In 2003, Australian botanist M. Clements proposed dividing Dendrobium between numerous segregate genera, noting (among other reasons) that the genus as previously recognised was not monophyletic. However, Clements' system does not seem to have garnered widespread usage with other orchid systematists preferring to retain a broad concept of Dendrobium (excluding some of the more egregious outliers) that largely corresponds with its established usage (e.g. Schuiteman 2011). Nevertheless, many of the subdivisions promoted by Clements remain recognised as well delimited groups. One such cluster is the assemblage of species recognised as Dendrobium section Aporum.

Growth habit of Dendrobium sect. Aporum, copyright Tony Rodd.

Species of section Aporum are epiphytes found in lowland forests of south-east Asia, extending eastwards to New Guinea and the Solomon Islands. Members of this section have thin stems that are erect at first but tend to become pendulous as they lengthen. Leaves are fleshy and equitant: that is, they are folded longitudinally with what would otherwise be the two sides of the dorsal surface fused, except at the base where they overlap with opposing leaves. The stem may be more or less completely concealed by the leaf bases. Tips of the leaves end in a point. Flowers are borne singly or in clusters, arising laterally on the stem between leaf nodes or at the tip of the stem alongside a terminal scale. The flowers may be subtended by persistent chaffy bracts. They are generally small and fleshy and tend to be short-lived, wilting after just a few days.

Flowers of Dendrobium anceps, copyright Aqiao HQ.

The functional significance of the Aporum section's distinctive leaves remains uncertain. As noted by Carlsward et al. (1997), the fleshy leaves might be taken as an adaptation to water retention. However, though access to water is a consistent concern for epiphytes, the humid rainforests in which Aporum species are found hardly seem the driest of places. Conversely, the effective even distribution of stomata on both sides of leaf resulting from their equitant condition may make it easier for excess water to be released from the plant.

Dendrobium distichum, photographed by Ronny Boos.

Orchids in general are, of course, most often considered by people as ornamental plants. My impression is that the various Aporum species tend not to be among the most widely grown of species though their unusual growth habit might attract interest. This may be due to them not being the easiest of orchids to maintain; they appear to require high humidity and warm temperatures to thrive with a cooler, drier period in the non-growing season. Among the more popular species are Dendrobium anceps and D. keithii, both of which produce small greenish flowers. Those of D. anceps have been described as having a distinct "apple pie" fragrance. Of course, if you happen to be wandering through the jungles of south-east Asia, you might well discover these plants growing of their own accord.

REFERENCES

Carlsward, B. S., W. L. Stern, W. S. Judd & T. W. Lucansky. 1997. Comparative leaf anatomy and systematics in Dendrobium, sections Aporum and Rhizobium (Orchidaceae). International Journal of Plant Sciences 158 (3): 332–342.

Clements, M. A. 2003. Molecular phylogenetic systematics in the Dendrobiinae (Orchidaceae), with emphasis on Dendrobium section Pedilonum. Telopea 10 (1): 247–298.

Schuiteman, A. 2011. Dendrobium (Orchidaceae): to split or not to split? Gardens' Bulletin Singapore 63 (1–2): 245–257.

The Huenellidae

Researchers who deal with the modern marine fauna are used to thinking of brachiopods as a marginal group, their diversity greatly overshadowed on a global scale by the superficially similar bivalves. However, modern brachiopods are but a shadow of their former selves; for much of the Palaeozoic era, their relationship with the bivalves was the inverse of today. Many are the brachiopod lineages that came and went over this time.

External views of ventral (left) and dorsal valves of Huenella triplicata, from Walcott (1924).

The Huenellidae were an assemblage of brachiopods that lived during the late Cambrian and early Ordovician (Amsden & Biernat 1965). They represent early representatives of the Pentamerida, a Palaeozoic order of fairly generalised-looking brachiopods. Within the Pentamerida, they fall within the suborder Syntrophiidina. Syntrophiidinans as a whole are rarely found in the fossil record and as a result remain poorly known. Members of the suborder share a distinctive shape with biconvex valves marked by a dorsal fold and ventral sulcus. That is, the midline of the shell is raised above either side with the ventral valve forming a 'valley' to match the raised 'hill' of the dorsal valve. What, if anything, was the purpose of this arrangement I wouldn't know but modern brachiopods often inhabit locations with a lot of organic silt and/or fine sediment. Perhaps the uneven level of the syntrophiidinan shell helped protect it from burial by a shifting substrate.

Interior view of ventral valve of Radkeina taylori, from Laurie (1997), with scoop-shaped spondylium at upper midline.

Families of Syntrophiidina may be distinguished based on the development of the spondylium, an internal projection at the base of the ventral valve that provided an attachment site for the shell muscles. Members of the Huenellidae possessed either a sessile spondylium or a pseudospondylium, a spondylium-type structure rising from the internal surface of the valve itself rather than from the hinge. Amsden & Biernat (1965) recognised a division of the huenellids between two subfamilies based on the development of the brachiophore plates, projections on the inside of the dorsal valve that would have supported the lophophore. Members of the Huenellinae possessed more developed brachiophores than members of the Mesonomiinae. Outer ornament of the huenellid shell varied from more or less smooth with weak concentric ridges to costate with distinct radiating ridges.

Phylogenetic relationships within the Syntrophiidina do not seem to have been established in detail but the early appearance in the fossil record of huenellids at least raises the question of whether they included the ancestors of later families. As well as other families of the Syntrophiidina, candidates for descent would include members of the suborder Pentameridina as well as of the related order Rhynchonellida. This latter order includes species which survive to the present day so the possibility exists that while the huenellids themselves may be long gone, their legacy may yet live on.

REFERENCE

Amsden, T. W., & G. Biernat. 1965. Pentamerida. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt H. Brachiopoda vol. 2 pp. H523–H552. The Geological Society of America, Inc.: Boulder (Colorado), and The University of Kansas Press: Lawrence (Kansas).