Caloppiidae

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

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

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

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

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

REFERENCE

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

Oribatid Time Again

The oribatid mite genus Neogymnobates was first recognised from Illinois in 1917. Since then, the genus has been found to be more widespread in North America and has also been described from Korea and Tibet. Species of Neogymnobates are known from arboreal habitats or in association with fallen wood, and live as grazers of micro-vegetation such as lichens.

Neogymnobates luteus, copyright Monica Young.

Neogymnobates belongs to the Ceratozetidae, a diverse family of oribatids whose characteristic features include a tutorium (a projecting tooth-like structure) on the side of the prodorsum and immovable pteromorphs on either side of the front of the notogaster. Neogymnobates has the lamellae on either side of the prodorsum widely separated from each other and connected by a transverse translamella at the front. There are thirteen pairs of setae on the notogaster and four pairs of porose areas (Balogh & Balogh 1992). One species, N. marilynae of British Columbia and Washington State, is known to have an extra unpaired porose area on the midline near the rear of the notogaster (Behan-Pelletier 2000), an unusual feature among oribatids but one whose significance is uncertain). Their legs end in three claws, a feature that (as I've commented before) correlates with their arboreal habits.

Half a dozen species of Neogymnobates have been recognised to date (Subías 2004). The species are distinguished by features such as the size and appearance of the setae, and the development of the prodorsal lamellae and translamella. One Korean species, N. parvisetiger, has been awarded its own subgenus Koreozetes due to its particularly small, almost indiscernable notogastral setae and its anteriorly notched rather than rounded rostrum (Aoki 1974). Most species are only known from limited ranges except one, N. luteus, for which separate subspecies have been recognised in northern North America and in Korea. Rather unexpectedly, this last species has also recently been recorded from Zanzibar (Ermilov & Khaustov 2018). This is a remarkable range increase, both geographically and ecologically (enough so that I can't help feeling it would benefit from double-checking) that raises the possibility that we may yet have a lot to learn about this oribatid genus.

REFERENCES

Aoki, J. 1974. Oribatid mites from Korea. I. Acta Zoologica Academiae Scientiarum Hungaricae 20 (3–4): 233–241.

Balogh, J., & P. Balogh. 1992. The Oribatid Mites Genera of the World vol. 1. Hungarian Natural History Museum: Budapest.

Behan-Pelletier, V. M. 2000. Ceratozetidae (Acari: Oribatida) of arboreal habitats. Canadian Entomologist 132: 153–182.

Ermilov, S. G., & A. A. Khaustov. 2018. A contribution to the knowledge of oribatid mites (Acari, Oribatida) of Zanzibar. Acarina 26 (2): 151–159.

Subías, L. S. 2004. Listado sistemático, sinonímico y biogeográfico de los ácaros oribátidos (Acariformes, Oribatida) del mundo (1758–2002). Graellsia 60 (número extraordinario): 3–305.

The Mites of the Incas

Diagnostic views of Haplozetes similis, from Gil & Subías (1993) as 'Lauritzenia (Incabates) sinuatus'.

The oribatid mite genus Incabates was first established by Marie Hammer in 1961 for a species from Peru (not surprisingly, with that name). Since then, species have been assigned to Incabates from tropical and subtropical regions almost throughout the world (though not, as yet, from the Ethiopian bioregion—Subías 2004). Incabates belongs to the Haplozetidae, an oribatid family distinguished by their possession of well-developed pteromorphs that are often, though not always, mobile, and jointed chelate-dentate chelicerae (Norton & Behan-Pelletier 2009; the nature of oribatid 'pteromorphs' has been explained in an earlier post). Incabates has been distinguished from other haplozetid genera by having a series of dorsal glandular openings on the body developed as tubular saccules, ten pairs of setae on the notogaster (the dorsum of the main body), four pairs of setae around the genital opening, and three claws at the end of each leg (Gil & Subías 1993; Weigmann & Monson 2004). The presence of three claws rather than one claw on each leg appears to be correlated with an arboreal lifestyle in oribatids (Karasawa & Hijii 2008), and Incabates species do appear to be mostly associated with forest habitats.

There is a lot of taxonomic instability within (and indeed, around) the Haplozetidae, and it remains uncertain at this point whether Incabates should be recognised as a distinct taxon. Genera of Haplozetidae have often been distinguished by combinations of characters, often simply numerical (such as numbers of setae) and it may be debatable to what extent these characters reflect actual relationships. The checklist of the world oribatid fauna by Subías (2004) lists Incabates as a subgenus of Lauritzenia. Gil & Subías (1993) separated Lauritzenia (including Incabates) from the related genus Haplozetes by the number of genital setae (four pairs in Lauritzenia vs five in Haplozetes) and then divided each of these genera into subgenera on the basis of claw number (tridactyl Incabates vs monodactyl Lauritzenia subgenus Lauritzenia). However, Weigmann (2010) argued that these characters might be too plastic to warrant generic distinction, and supported treating all as a single undivided genus Haplozetes until the relationships within the group were better established. As yet, that's something we're still waiting on.

REFERENCES

Gil, J., & L. S. Subías. 1993. La familia Haplozetidae Grandjean, 1936 (Acari, Oribatida) en la Península Ibérica. Mediterránea Ser. Biol. 14: 23-30.

Karasawa, S., & N. Hijii. 2008. Vertical stratification of oribatid (Acari: Oribatida) communities in relation to their morphological and life-history traits and tree structures in a subtropical forest in Japan. Ecological Research 23 (1): 57-69.

Norton, R. A., & V. M. Behan-Pelletier. 2009. Suborder Oribatida. In: Krantz, G. W., & D. E. Walter (eds) A Manual of Acarology, 3rd ed., pp. 430-564. Texas Tech University Press.

Subías, L. S. 2004. Listado sistemático, sinonímico y biogeográfico de los ácaros oribátidos (Acariformes, Oribatida) del mundo (1758-2002). Graellsia 60 (número extraordinario): 3-305.

Weigmann, G. 2010. Oribatid mites (Acari: Oribatida) from the coastal region of Portugal. IV. The genera Coronoquadroppia, Scheloribates, Haplozetes and Pilobates. Soil Organisms 82 (3): 383-406.

Weigmann, G., & F. D. Monson. 2004. A new genus and species of Haplozetidae (Arachnida: Acari) from Great Britain with a key to the European genera. Journal of Natural History 38 (11): 1415-1420.

On a Wing and a Mite

Dorsal and ventral views of Monogalumnella neotricha, with legs removed. Figures from Balogh & Balogh (1992).

In a previous post, I introduced you all to the oribatid mites. Oribatids come in a wide range of varieties, and the animal in the figures above is a member of the oribatid family Galumnellidae. Galumnellids belong to a group of oribatids, the galumnoids, marked by their well-developed pteromorphs: the roughly triangular structures at either side of the front of the body. Many oribatids have pteromorphs developed to a greater or lesser degree, but the pteromorphs of galumnoids are particularly noteworthy for their size and for the development of a hinge between the pteromorph and the main body, so that the pteromorph can be folded down to cover the legs for protection (other species have the pteromorphs as fixed outgrowths of the body). The name 'pteromorph', of course, means 'wing-shaped', and you can readily find cases where galumnoids have been referred to as 'winged mites' (especially in older publications). Woodring (1962) even suggested that galumnoids might provide a useful analogy for the evolution of wings in insects. However, pteromorphs are not actually wings like those of insects, being used only for protection, not flight. In animals as small as oribatids, the relative viscosity of the air becomes very high, not to mention the relative force of small air movements. Vary small arthropods that move aerially either develop long hairs or similar structures so that they can be passively lifted and carried by the breeze (like the line of silk produced by ballooning spiders) or have reduced wings with long fringes of hairs to maintain wing surface area while minimising air resistance (such as mymarid wasps, thrips or ptiliid beetles). A solid plate like the galumnoid pteromorph would be to difficult to move*.

*Similar issues affect suggestions that the absent fossil record of the earliest winged insects may indicate that flight evolved at small sizes. It seems almost certain that the first flying insects were relatively large.

The Galumnellidae can be distinguished from other galumnoid mites by the lack of protruding lamellae on the prodorsum (the top of the 'head'), the pointed rather than rounded rostrum, and the shape of their chelicerae. The chelicerae of galumnellids are long and slender, compared to the shorter, stronger chelicerae of their relatives in the Galumnidae. Galumnella has been shown in the laboratory to be panphytophagous (Badejo & Akinwole 2007)—that is, it will accept any type of plant or algal food, both living and dead.

REFERENCES

Badejo, M. A., & P. O. Akinwole. 2007. Preliminary study of the feeding habits of seven species of oribatid mites from Nigeria. Systematic and Applied Acarology 12: 121-125.

Balogh, J., & P. Balogh. 1992. The Oribatid Mites Genera of the World, 2 vols. Hungarian Natural History Museum: Budapest.

Woodring, J. P. 1962. Oribatid (Acari) pteromorphs, pterogasterine phylogeny, and evolution of wings. Annals of the Entomological Society of America 55 (4): 394-403.