A Catalogue of South(ern) African Insects
BIOSIS Internet Resource Guide - Collembola
Gordon Ramel's Springtail Page
The Collembola page of The Tree of Life
Catalogue of the Collembola of the World
Botany and Zoology, Australian National University, GPO Box, ACT 0200, Australia
© Copyright of this page belongs to the author.
Worldwide, nearly 7,500 collembolan species in 581 genera have been described. In South Africa, 69 genera and subgenera, and 152 species, belonging to 13 families are known, with a further 16 species inquirenda. These are species that have been identified, but probably incorrectly so. It is clear that only a small proportion of the fauna has been described. The most recent English general text on Collembola is that of Hopkins (1997).
There are three suborders of Collembola which can be easily distinguished on body shape. The Arthropleona, to which most species belong are elongate, with most body segments separate and equal in length. The Symphypleona are globular with anterior body segments fused and the Neelipleona are a very small group of minute globular animals with live deep in the soil. There are about 25 families with most families in the Arthropleona.
When such a small proportion of the fauna is known from the region, there is no sound basis from which to make comments on the characteristics of the fauna. A comparison of the genera with those from other southern continents indicates that the soil fauna of southern Africa is not markedly different in composition to that of other continents. In the Neanurinae (Neanuridae), there is an indication that the genus Ectonura may have radiated on the continent. The Neanurinae are associated with rotting wood especially sticks. The epigaeic fauna, such as is found on grasses, shows the most differences from other continents in that the genus Seira (Entomobryidae) is very diverse with 22 species. In Australia, for instance, with vast expanses of savannah woodland similar to those in southern Africa, there is only one species of Seira known from this habitat. Similarly in the Symphypleona (Bourletiellidae), the dominant genus found on grasses is Prorastriopes while in Australia it is Corynephoria, a genus not found in southern Africa. Prorastriopes does occur in Australia but native species found only in the north of the continent.
The endemism of the fauna at species level may be high, but most genera recorded are cosmopolitan in distribution. Among the 69 genera recorded, only ten (ca 15%) are currently endemic to southern Africa. These are Odontellina, Najtafrica, Probrachystomellides, Orthonychiurus, Fissuraphorura, Caprobrya, Lepidodrugeeria, Calobatinus, Pseudocyphoderus, Neophorella and Triosminthurus.
Collembola are not well known because of their small size and cryptic habits. The order as a whole is, however, extremely widespread with a global distribution that includes polar and arid regions and is particularly diverse in tropical regions. Although Collembola are predominantly soil and litter dwellers they also occur in a wide range of other habitats such as on vegetation including tree canopies, in caves, in the marine littoral zone and in freshwater systems. Average densities in soils are usually between 10,000 and 30,000 per m2 but can be as high as a million in rare situations and so Collembola may outnumber mites that are usually the most abundant micro arthropods. As detritivores, Collembola are an important group in nutrient cycling and, can be considered beneficial organisms. Very few species feed on live plant material and are considered pests. The biology and widespread nature of Collembola suggest that they warrant more attention from biologists.
Springtails, the common name is derived from the ability of many species to leap considerable distances when disturbed, are soft-bodied arthropods. They are allied to insects and often included in the Class Insecta as they are similar in having a body divided into three parts: a head with antennae and two pairs of mouthparts, a three- segmented thorax with a pair of jointed legs on each segment, and an abdomen with paired appendages ventrally on some segments. However they differ from insects in lacking a hard exoskeleton and wings, in the mouthparts being internal and in possessing simple eyes, maximally 8, on each side. Consequently, Collembola are occasionally erroneously referred to inaccurately as "primitive insects". Adult animals are small compared to most insects most species being only 0.5-3 mm long, but a few are as long as 10 mm or as short as 0.2 mm as adults. Many species are white or black or various shades of grey but others are brightly coloured and attractively patterned. Collembola are furnished with chaetae (setae), which can be numerous or sparse, fine or thick, long or short, serrated, ciliated, clavate or smooth. Scales and bothriotricha (long, usually ciliated, slender chaetae inserted in pits) are found in some families.
The cuticle of Collembola is composed of triangular granules that may be fused into larger raised tubercles, rendering it hydrophobic. The hydrophobic nature of the cuticle enables the animals to float on water. Most species are terrestrial or arboreal but a few groups are considered aquatic. Some aquatic species live exclusively on water surfaces but others live more intimately with the aquatic environment: interstitially in sand and under submerged stones and rocks in streams and on the sea shore. It has been reported that if collembolan eggs hatch underwater, the juveniles and even adults can exist totally submerged. However if their cuticle becomes exposed to air, structural changes take place making it repel water and prevent further submersion. Most limnologists encounter Collembola most frequently, and often abundantly, in drift nets when they have been washed from surrounding terrestrial habitats into streams where they are able to float. If numerous, they may form rafts of live animals. Running water is considered a major conduit for dispersal to new habitats; consequently, Collembola might be considered as facultatively surface-aquatic animals for at least some part of their life histories. Some authors (e.g. Waltz and McCafferty, 1979) have attempted to classify Collembola according to the degree to which they utilise aquatic environments. As can be seen from the above description of the ecology of the group, such attempts are simplistic in that they are dividing what is essentially a continuum and obscuring subtleties of ecological variation.
Collembola are small, wingless hexapods, either pigmented or white, normally 1-2 mm long; the mouth-parts are enclosed within the buccal cavity (mouth); one pair of elongate maxillae and one pair of mandibles are enclosed by the labrum and labium. A post-antennal organ is usually present; there are usually up to eight ocelli on each side of the head, but sometimes fewer ocelli are present the word these is one to be avoided or even totally absent; the antennae are four-segmented, with muscles within all segments the antennal segments are sometimes sub segmented; the thorax has three segments, the first segment being sometimes lacking chaetae and shorter than the other segments; the legs and the abdomen are both six-segmented but the segments in the abdomen are frequently fused; specialised appendages, the ventral tube, tenaculum and furca, are found ventrally on abdominal segments I, III; the genital opening is in a ventral position on abdominal segment V; the anal aperture is on abdominal segment VI; cerci are absent, but anal spines are sometimes present; metamorphosis absent.
MORPHOLOGY
The Head:
The mouth is anterior and the head directed along the main body axis, except in Symphypleona and Neelipleona where the head is at right angles to the body; the antennae have four-segments which may be subdivided into at least two, but sometimes numerous, sub segments the antennae vary in length from being shorter than the head to being longer than the body and are sometimes modified in the male to form a clasping organ. There are a maximum of 8 + 8 ocelli on pigmented patches on each side of the head. Mouthparts are elongate and adapted either for biting and chewing/grinding, or for fluid feeding. They consist of a pair of maxillae, a median labium and a median labrum. A mandible usually present, frequently with finely-toothed molar plate, the head nearly always with one or more teeth; maxillary head with three or four teeth at most, and characterised by the presence of a maximum of six fringed lamellae and a pair of posterior extensions (fulcra) on which the cardo articulates. The pleural folds, together with the labrum and the labium, completely enclose the mouthparts to form the buccal cone, which may project anteriorly.
The Thorax.
The thoracic segments are conspicuous in the Arthropleona but not in Symphypleona or Neelipleona where they are fused and form a single mass with the abdomen; thoracic segment 1 is always reduced.
The Abdomen.
The abdomen is six-segmented, but segments I-IV, I-V, IV-VI, V-VI or IV-V may be fused dorsally; segment I carries a ventral tube, segment III a hook or tenaculum and segment IV a furca or 'spring'. The ventral tube consists of a column with a pair of tubes at the end, which are normally retracted within the column, but which can be extruded to absorb moisture or provide adhesion to the substrate after a leap, the 'jumping organ' or furca is normally folded forward beneath the body and held in place by the teeth of the tenaculum. When suddenly released from the tenaculum, it flexes backwards to the resting position and so hits the ground with considerable force, propelling the animal into the air. Jumping is both a reaction to disturbance and a means of dispersal and some directional ability has been demonstrated. The genital aperture is positioned ventrally on abdomen V and consists of a transverse slit in the female and a raised and longitudinal slit in the male; both supplied with either a few, or with numerous short chaetae. The anus is located on abdomen VI.
Immature Stages
Eggs are spherical and pale. Juveniles resemble adults and it is sometimes difficult to distinguish either on size or on morphology, although complete adult chaetotaxy and colour gradually develop during the pre-adult instars. Differences seen between 1st and 2nd instars and the last preadult instar and adult, are insufficient to be classified as metamorphosis. There are normally 3 - 13 stages before maturity but four or five stages are also common. The life cycle from egg to adult can take on average from one to three months but this can vary in different species and at different temperatures. Individuals of some species may live for five years, yet in other species, males mate with preadult females and then die, living only a few days in total. Adults continue moulting throughout life, and may undergo up to 60 moults in some species, although no further increase in size usually occurs after about moult 15.
Upton (1991) is a recent general textbook on collecting and preserving insects, which has relevance to Collembola.
a) Hand collections. This is the only efficient method of collecting some species that live in cryptic habitats and do not emerge readily or have low density populations Searching can be carried out by looking under rocks and stones in and at the edge of water bodies, from the surface of water in puddles, and other water bodies. Material can be tipped onto a white tray, white plastic sheet or plate and specimens aspirated into a tube in the usual way with a pooter or aspirator. A fine paintbrush can be used to brush animals from beneath stones that have been submerged in water on a tray or the stone tapped on the tray. Surface aquatic species are collected with a fine-meshed aquarium net.
b) Pitfall traps. Pitfall traps are successful in collecting aquatic species if used at the edges of water bodies, both still and running, and below high tide on beaches. McCartney bottles (1-2 cm external diameter and 5-8 cm depth), three quarters filled with 90-100% ethyl alcohol), are buried into the soil so that the rim of the bottle is flush with the soil surface. Other preservative may be used, as ethyl alcohol is not neutral, but attractive for some Collembola. It is easiest to clear an area of about 15 cm diameter around each trap as this improves catch and visibility. If the weather is dry, a few drops of glycerol should be added to the alcohol in the traps to retard evaporation. After 4 to 5 days the bottles are removed, topped up with 96% ethyl alcohol, labelled and capped.
Collecting from river banks, sand banks and beaches. On coastal sand dunes as well as river banks, a hole can be dug in the sand to a depth of about 30 cm and water poured into it for several days, the damp sand dug up and Collembola removed from it by funnel extraction or flotation. Alternatively, on beaches at least a metre or so above the water edge, a hole can be dug down to the water table. In this case Collembola fall from the sides of the hole onto the water surface where they float and can be retrieved using a fine aquarium net.
Killing and preserving
Collembola are generally considered to be insects, but because of the differences they exhibit from insects, i.e. soft bodies, fragile appendages, wide range of colours and patterns and easily detached surface structures, rather different methods must be used for their preservation.
The fluid usually recommended for both killing and preservation Collembola is 95% or absolute ethanol. It is essential to ensure the Collembola are totally submerged immediately in the preserving fluid. Floating specimens should be decanted, more alcohol of high concentration added and the tube capped, inverted and then vigorously shaken as soon as possible after collection. This encourages the specimens to sink but if they are left floating for long or remain in low concentrations of alcohol for any length of time, it is harder to make them sink. Specimens do become brittle and difficult to mount with age and chaetae loose their refractory properties after long storage in alcohol.
Preparation of specimens for detailed study
Collembola, being small, soft-bodied invertebrates, present different and greater problems for examination than do most other arthropods. They may be mounted on slides using the following method:
Specimens are left in lactic acid until clear which may take from a few minutes to several days depending on the specimen. Heating the solution for a few seconds may be necessary with deeply pigmented or old specimens. A drop of the gum arabic Berlese fluid is then placed on a slide and the specimen placed in it. Round cover slips, no. 0 or 00, of 10 mm diameter are easiest to use, but are much more expensive than square cover slips.
Life history
The life histories of only a few Collembola have been studied in detail anywhere in the world and there is no information at all on South African forms in this respect. In general, the eggs of Collembola are laid singly or in clusters in protected sites such as in soil, in leaf litter, under stones or in crevices. In some Symphypleona, eggs are covered with freshly-eaten soil mixed with a rectal fluid voided through the anus after oviposition. This covering protects the eggs against dehydration and fungal attack. In other species, eggs are kept free of fungal hyphae by the grazing of adults. The time for hatching and development varies with species and with temperature. Instars are similar to the adult in general appearance, the only difference being smaller size, lack of genital apparatus and lack of development of the full chaetotaxy.
Distribution and habitats
Terrestrial Collembola are found in a wide variety of usually moist habitats, predominantly in soil and leaf litter and other decomposing materials such as logs and dung. Certain species are found on grasses, in flowers, under the bark of trees or in tree canopies. Collembola are virtually ubiquitous, being found in all terrestrial biomes, including mountain tops, Polar Regions and deserts. Many species inhabit caves.
Some species seem to be distributed readily in surface run-off. Some factors influencing the distribution of interstitial Collembola are the location of food, disturbance, and soil pore space. Collembola generally are sensitive indicators of disturbances, such as oil pollution on beaches and chemical pollution of rivers and streams and therefore are of value in environmental assessment.
Semi-aquatic and facultatively aquatic species are common in marine and freshwater littoral habitats, occurring interstitially in wet sand, on and under smooth stones and roughly surfaced rocks including coral reefs, and on water surfaces. Only a small number of species can be considered strictly aquatic and live on water surfaces throughout life. They belong to Sminthurides and allied genera in the Sminthurididae and live on the surfaces of fresh, still waters.
A larger number of taxa are considered marine littoral or facultatively aquatic. They occur on riverbanks and in the intertidal zone occupying the interstitial soil and sand habitat and are also found under stones and plant debris and in fissures in rocks. Oudemansia and Archistoma, genera that are exclusively marine littoral, have been recorded from southern Africa. Ballistura, a genus restricted to habitats adjacent to fresh water, has also been recorded from southern Africa.
Feeding behaviour
Most species feed on micro-organisms associated either with the water surface, the rhizosphere, with decomposing organic matter, with fungal fruiting bodies or, to a lesser extent, with the aerial surfaces of plants; they do not feed readily on sterile leaves. Collembola act as catalysts in the breakdown of organic matter and in the cycling of plant nutrients by grazing on and distributing propagules of microorganisms and, through their feeding and other activities and the deposition of faecal material, they can alter the physical properties and structure of soils.
Reproduction
Sperm transfer is indirect in most families, the males depositing a globular, stalked spermatophore on the ground in which sperm is encysted. In some Symphypleona (Bourletiellidae and Sminthurididae) there may be an elaborate dance, in which the male grasps the female by means of the specialised spines and hairs on the antennae, head or legs and directs her to the spermatophore. There is strong sexual dimorphism in these families and all truly aquatic species belong to this group.
Predators
Many predacious arthropods prey on Collembola as do coral reef fish, wading birds, small mammals, reptiles and frogs. The predatory arthropods, some of which have evolved elaborate catching devices, include carabid and staphylinid beetles, dacetine ants, Hemiptera, empidid and dolichopodid flies, spiders, opiliones, pseudoscorpions, centipedes, and prostigmatid and mesostigmatid mites. For instance they comprise the main items of diet for bdellid, cunaxid and anystid mites. However some families, notably the Onychiuridae and Neanuridae, seem immune from predators, probably because they contain and sometimes exude, a toxic or distasteful chemical. Viruses, bacteria, fungi and Protozoa as well as nematodes have been found to be parasites of Collembola. Apart from leaping and cryptic coloration, Collembola have a number of defence mechanisms against predation. These include immobility, mimicry and a spiny dorsum.
Please refer to the references listed below, which is reasonably complete up to 1980. For more information, visit
A Catalogue of South(ern) African Insects
BIOSIS Internet Resource Guide - Collembola
Gordon Ramel's Springtail Page
The Collembola page of The Tree of Life
Catalogue of the Collembola of the World
J. A. Barra compiled this list based on bibliographical data and list of species from South Africa only found on the Rhodes University website. It includes also the described species from the countries of southern Africa, such as Namibia, Botswana, Zimbabwe, Mozambique and the new Provinces of South Africa. As the taxonomic identity of many of the species is uncertain, only numbers of species in each genus are given and not species names.
Members of this family are terrestrial, found in numerous habitats (soil, under bark, seashore, mushroom beds).
| Acherontiella | Absolon, 1913 | 1 sp. |
| Ceratophysella | Börner, 1932 | 3 spp |
| Choreutinula | Paclt, 1944 | 1 sp. |
| Hypogastrura | Bourlet, 1839 | 4 spp |
| Mesogastrura | Bonet, 1930 | 1 sp. |
| Willemia | Börner, 1901 | 2 spp |
| Xenylla | Tullberg, 1869 | 7 spp |
The members are terrestrial.
| Odontella | Schäffer, 1897 | 1 sp. |
| Superodontella | Stach, 1949 | 1 sp. |
| Odontellina | Deharveng, 1981 | 1 sp. |
Most are terrestrial, found in various habitats (soil, leaf litter, under bark and stones, littoral habitats).
| Frieseinae | ||
| Friesea | Dalla Torre, 1895 | 3 spp |
| Pseudachorutinae | ||
| Aethiopella | Handschin, 1942 | 2 spp |
| Anurida | Laboulbène, 1865 | 1 sp. |
| Najtafrica | Barra (in press) | 1 sp. |
| Oudemansia | Schött, 1893 | 1 sp. |
| Pseudachorutella | Stach, 1949 | 1 sp. |
| Pseudachorutes | Tullberg, 1871 | 2 spp |
| Neanurinae | ||
| Ectonura | Cassagnau, 1980 | 3 spp |
| Neanura | MacGilliway, 1893 | 1 sp. |
| Vitronura | Yosii, 1969 | 1 sp. |
Most brachystomellids are confined to damp soil.
| Brachystomella | Agren, 1903 | 3 spp |
| Probrachystomellides | Weiner and Najt, 1991 | 1 sp. |
| Setanodosa | Salmon, 1942 | 1 sp. |
Members of this family are leaf litter or soil-dwelling forms.
| Onychiurinae | ||
| Onychiurus | Gervais, 1841 | 1 sp. |
| Orthonychiurus | Stach,1954 | 1 sp. |
| Protaphorura | Absolon, 1901 | 1 sp. |
| Tullbergiinae | ||
| Fissuraphorura | Rusek, 1991 | 1 sp. |
| Mesaphorura | Börner, 1901 | 2 spp |
| Paratullbergia | Womersley, 1930 | 1 sp. |
| Tullbergia | Lubbock, 1876 | 2 spp |
Most species live in soil, but also associated with freshwater or in crevices on the seashore.
| Archisotoma | Linnaniemi, 1912 | 2 spp |
| Arlea | Womersley, 1939 | 1 sp. |
| Ballistura | Börner, 1906 | 1 sp. |
| Clavisotoma | Ellis, 1970 | 1 sp. |
| Cryptopygus | Willem, 1901 | 5 spp |
| Folsomides | Stach, 1922 | 3 spp |
| Folsomina | Denis, 1931 | 1 sp. |
| Isotomiella | Bagnall, 1939 | 4 spp |
| Isotomodes | Linnaniemi, 1907 | 1 sp. |
| Isotomurus | Börner, 1903 | 5 spp |
| Micranurophorus | Bernard, 1977 | 1 sp. |
| Parisotoma | Bagnall, 1940 | 1 sp. |
| Proisotoma | Börner, 1901 | 2 spp |
Terrestrial. Members are found in a wide range of habitats (leaf litter, on the soil, under bark, in caves).
| Capbrya | Barra,1999 | 2 spp |
| Coecobrya | Yosii, 1956 | 1 sp. |
| Entomobrya | Rondani, 1861 | 5 spp |
| Lepidocyrtus | Bourlet,1839 | 4 spp |
| Lepidokrugeria | Coates, 1969 | 1 sp. |
| Pseudosinella | Schäffer, 1897 | 4 spp |
| Seira | Lubbock, 1869 | 22 spp |
Terrestrial. They are found on trees, grasses, superficial layers of the litter, some live in caves.
| Paronella | Schött, 1893 | 1 sp. |
Terrestrial. All members occur in the nest of ants and termites.
| Calobatinus | Silvestri, 1918 | 1 sp. |
| Cyphoda | Delamare, 1948 | 3 spp |
| Cyphoderus | Nicolet, 1842 | 5 spp |
| Pseudocyphoderus | Imms, 1912 | 1 sp. |
Terrestrial. Members are found on soil surface, leaf litter, some species live in caves.
| Neophorella | Womersley, 1934 | 1 sp. |
Terrestrial. All species live in soil or in caves.
| Megalothorax | Willem, 1900 | 2 spp |
The following families are terrestrial. They live predominantly in superficial leaf litter, on low vegetation, on trees, on surface of still fresh water.
| Denisiella | Folsom and Mills, 1938 | 1 sp. |
| Sphaeridia | Linnaniemi, 1912 | 2 spp |
| Katianna | Börner, 1906 | 1 sp. |
| Sminthurinus | Börner, 1901 | 3 spp |
| Stenognathellus | Stach, 1956 | 1 sp. |
| Bourletiella | Banks, 1899 | 1 sp. |
| Prorastriopes | Delamare, 1947 | 4 spp |
| Rastriopes | Börner, 1906 | 1 sp. |
| Tritosminthurus | Snider, 1988 | 1 sp. |
| Papirinus | Yosii, 1954 | 1 sp. |
| Sminthurus | Latreille, 1802 | 1 sp. |
Terrestrial. Species are found on surface soil, on the bark of trees.
| Dicyrtomina | Börner, 1903 | 2 spp |
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