African Invertebrates Vol. 56 (3): 527– 548 Pietermaritzburg 25 November 2015
Eco-taxonomic prole of an iconic vermicomposter — the ‘African
Nightcrawler’ earthworm, Eudrilus eugeniae (Kinberg, 1867)
Robert J. Blakemore
VermEcology, Yokohama, Japan and
C/- Biodiversity Lab., College of Natural Science, Hanyang University, Seoul 133-791;
Eudrilus eugeniae (Kinberg, 1867), an adaptable exemplar of an anatomically advanced earthworm having
direct fertilisation, is reviewed ecologically. A tropical West African species originating in savannah soils, it
thrives on organically rich substrates. It has a rapid life-cycle from cocoon to maturity in as little as 47 days.
Presence of this worm raised experimental pasture yields up to 83.9 %. Copious pellet-like casts deposited
onto the soil surface are sought by roots. Passage of organic material through its gut reduces microbial
pathogens and the resulting vermicompost product has enhanced nutrients, and microbial and enzymatic
properties. Preliminary pharmaceutical reports are of inhibition of ‘Golden staph’ Staphylococcus aureus and
‘Thrush’ Candida albicans, plus anti-tumour effects in cancer cell lines. Its handling characteristics make
or ‘seed’ cultures, (2) high-protein worm biomass for stock feeds, (3) organic fertiliser, (4) bio-stabilisation
matter (SOM, or humus), (7) bio-prospecting for pharmaceuticals, cosmetics or ‘silk’, and (8) eco-toxicology/
ethology research. New reports are of cultivation in Denmark, South Africa, Egypt, Saudi Arabia, Ecuador,
Peru, Indonesia, Malaysia, Thailand and Vietnam. Eudrilus eugeniae
global distribution and taxonomy updated with mtDNA barcodes.
KEY WORDS: Annelida, Oligochaeta, Eudrilidae, vermiculture, DNA barcoding, soil ecology, megadrile
This paper reviews the current knowledge of the eco-taxonomic and morpho-molecular
Eudrilus eugeniae (Kinberg, 1867). It is not known when its vermiculture
potential was initially recognised but its initial wide ‘expat’ distribution has been mainly
attributed to accidental human transportation, since it was already well established in
the early 1800s (e.g. Perrier 1872; Michaelsen 1900, 1903; Stephenson 1923). More
recently it has been deliberately introduced for commercial and experimental purposes.
Ecological studies commenced from the mid-1960s to the late 1980s (e.g. Eno 1966;
Madge 1969; M’ba 1978; Neuhauser et al. 1979; Graff 1981; Bano & Kale 1988). This
of Lee (1959) as reported by (Blakemore 2008b) since it responds facultatively, spanning
the spectrum as either a geophage ‘topsoil’ species or alternatively as a detritivore
‘litter’ species. Lee (1985) characterised it as a topsoil species and it is known to deposit
per annum according to Gates (1972: 52) — as well as producing uniformly enhanced
vermicomposts when reared on diverse organic ‘waste’ substrates.
eugeniae is also reviewed and supported with mtDNA COI barcodes (see Appendix).
a detailed species description. Further data is presented from comparative ecological
528 AFRICAN INVERTEBRATES, VOL. 56 (3), 2015
studies conducted by the author (Blakemore 1994, 1997, 2008a). This update notes
the confusion over reproductive and digestive organs, counters misdescriptions (such
as that by Vijaya et al.
distribution reports, such as New Zealand by Sims and Gerard (1985, 1999). A summary
of the glo bal distribution for Eudrilus eugeniae is updated from Michaelsen (1900,
1903). New ecological data is consolidated.
MATERIAL AND METHODS
Earthworms from worm farms in Australia and the Philippines were studied, and
and conventions of Blakemore (2012b). PCR methods similar to those described in
Blakemore et al. (2010) were used for mtDNA barcoding. Results of genetic analyses
with BLAST programs (www.blast.ncbi.nlm.nih.gov/BLAST.cgi) are compared to
Genbank (genbank.com) in the Appendix. Laboratory and glasshouse experiments plus
by Blakemore (1994, 1997, 2008a) with extensive literature searches; herein data is
integrated and compared with recent published reports. Abbreviations: ANC = African
Nightcrawler; np = nephropores; Qld = State of Queensland in tropical NE Australia;
rhs/lhs = right/left hand side.
Phylum Annelida Lamarck, 1802
Class Oligochaeta Grube, 1850
Order Megadrilacea Benham, 1890
Family Eudrilidae Claus, 1880
Eudrilus eugeniae (Kinberg, 1867)
Lumbricus eugeniae Kinberg, 1867: 98. [Type locality: Humid mounts and valley of St Helena (15°56'S
05°43'W). Types in Natural History Museum, London BMNH 1904.10.5.550 with Swedish
Museum label: “Lumbricus Eugeniae Kinberg St Helena Swed. State Museum.” The specimen
Eudrilus decipiens Perrier, 1871: 1176; 18721887: 247 (syn.: lacazii, peregrinus,
Eudrilus lacazii Perrier, 1872
Eudrilus peregrinus Perrier, 1872
Eudrilus roseus Michaelsen, 1892
2162. Michaelsen notes “?Eudrilus perigrinus
Eudrilus eugeniae: Beddard 1895lacazii, peregrinus, decipiens, boyeri, sylvicola, jullieni,
roseus); Eisen 1900 decipiens, lacazii +
peregrinus Perrier, 1872; boyeri, sylvicola, jullieni, roseus, erudiens); Stephenson 1923: 486;
1930: 873; Gates 1942
386; Csuzdi & Pavlicek 2009: 13 (excluding the peregrinus synonym by oversight?); Blakemore
1994; 2002; 2012b; 2013; 2014: 122.
Etymology: Named after Johan Gustaf Hjalmar Kinberg’s Swedish survey ship, the
BLAKEMORE: EUDRILUS EUGENIAE 529
Body length: Complete matures, 90–185 mm (pers. obs. and Gates 1972) or up to
250– 400 mm under optimal culture conditions (Viljoen & Reinecke 1994; Parthasarathi
1982). Width: Approximately 4–8 mm.
Mass: Mean per adult ca. 1.0 g (pers. obs.) or optimal maximum 5. 0–6.0 g.
Segments: 161–211 (pers. obs. and Gates 1972) or 250–300 (Viljoen & Reinecke 1994,
suggesting that larger worms add segments); constriction of 40– 46 seen in several Qld
specimens may be artefactual.
Colour: Red-brown dorsum fading posteriorly; anterior with bright blue/green iridescent
sheen from cuticle diffraction, ventrum beige, clitellum darker (sometimes lighter) than
Prostomium: Small, open epilobous.
Dorsal pores: None.
Setae: Eight per segment from 2, closely paired; setae a–b on 17 absent (dehisced); ratio
of aa:ab:bc:cd:dd:U on 7 = 6:1:5:1:10:0.5. Penial/genital setae absent.
Nephropores: Just behind anterior furrow of each segment (longitudinal slits) from 3/4
in c lines or slightly more median (sometimes in d lines).
Clitellum: 13, 14, 15– 18, usually 13, 14–18 and interrupted ventrally.
Male pores: In 17 on tips of longitudinally grooved, tapering, eversible penes in large
ventral chambers, retracted as lateral slits with wrinkled lips just anterior to 17/18 in
line with b setae.
in 14 as raised intrasegmental openings just anterior to c setae. Gates (1972: 51) calls
these “vaginal apertures”.
Genital markings: Central raised pad centred in 17 between male pores, faintly repeated
Fig. 1. A Eudrilus eugeniae specimen from a vermicompost site.
530 AFRICAN INVERTEBRATES, VOL. 56 (3), 2015
Beddard 1895 Eisen 1900
BLAKEMORE: EUDRILUS EUGENIAE 531
Septa: From 4/5; (6/)7/8/9 and 14/15 thickened.
Dorsal blood vessel: Single, truncated before anterior hearts in 7; according to Gates
(1972: 51) connects to paired supra-oesophageals in 7– 14 and paired extra-oesophageals
median to the hearts.
Hearts: In 7 lateral, in 8– 11 latero-oesphageal, all distended with blood in some Qld
specimens (cf. Gates (1972) who said the anterior hearts were undistended).
Gizzard: Weakly muscular in 5 immediately behind pharyngeal mass.
Calciferous glands: Ventral spheroidal sacs in 10 and 11 (concealed by seminal vesicles):
large and pink due to blood supply with many internal lamellae; also in 12 (concealed
by seminal vesicles) a pair of yellow, lobular ‘calciferous’ glands which are medially
placed lateral to the oesophagus and ducted posteriorly into it in 13. This latter pair
calls the median oesophageal sacs “chylustaschen” but Stephenson (1930) only called
the paired glands in 12 “calciferous”. Eisen (1900: 138) found neither crystals nor lime
granules in the paired “diverticles” in 12, whereas Gates (1972: 51), after claiming
calcareous granules in both median and paired glands, classed them all as calciferous.
Intestine: Origin in 14 or close to 14/15. Caeca and typhlosole absent. Small, supra-
intestinal glands present in eight to forty-two segments in some of 62– 132 (Gates 1972:
52; 1982: table 8) may assist digestion and/or be implicated in the immune competency
of the worms.
Nephridia: Paired, large coiled holonephridia in each segment from 4, not obviously
Male organs: Holandric with two large, unpaired (or attached?) sacs seen ventrally in 10
and 11, each contain a testis anteriorly and funnels posteriorly, i.e. two pairs of testes in
testes funnels are small and free from iridescent spermatozoa which aggregate in the
ducts and thus are easily missed. The male apparatus is complicated and descriptions
differ somewhat; the copulatory chamber contains a pointed and curved penis plus a
describe as a “Y-shaped gland” that opens into a groove going nearly to the tip of the
penis. Eisen found the product of this Y-shaped gland to be a secretion similar to that of
the silk gland of a caterpillar (possibly analogous to penial setae as found, for example,
in Nsukkadrilus mbae Segun, 1977, to remove sperm of previous concopulant?). The
Y-shaped gland is lacking in Eudrilus pallidus Michaelsen, 1891 and the copulatory
Fig. 2. Eudrilus eugeniae: (a) ventral view of Qld specimen, (b) vasa deferentia unite to form the muscular
euprostates ducting to the centre of the copulatory chamber (characteristic Y-shaped gland on rhs ducts
to lhs), (c) ‘spermathecal’ aperture and combined oviduct (unravelled) to ovisac opposite saccular
gland at junction of duct and ‘ampulla’ (ovary not shown), (d) prostomium, (e) calciferous glands,
hearts and dorsal vessel, (f) dorso-lateral view of caudal segments narrowing to pygomere, (g) cocoon.
sometimes fused to form a “single horseshoe-shaped
called the silk-producing “Y-shaped gland” (indicated as “Y-sg”).
532 AFRICAN INVERTEBRATES, VOL. 56 (3), 2015
chambers are absent from E. simplex Michaelsen, 1913, serving to anatomically separate
them from E. eugeniae according to Beddard (1895) and Gates (1972: 51).
calls this the ‘diverticulum’) or duct by long, coiled oviduct tubes in 14, sited opposite
a saccular gland. Eisen’s (1900: 139) description differed from Beddard’s (1895) but
both (mistakenly?) agreed that ovaries in 13 are combined with ovisacs; and, whereas
Eisen thought there were two pairs of ovaries in segment 13, Gates (1972: 52) had the
second, functional pair in 14. However, Michaelsen (1892
small ovaries paired behind septum 12/13 connecting with the saccular part of the
spermatheca (what Sims (1987) calls the “receptaculum seminis”) and that the ovisac
or “receptaculum ovorum” is terminal to a long second oviduct. Easily missed, this
oviduct usually connects with the larger oviduct leading to the ovisac where the eggs
mature (as described by Eisen 1900: 139). Histological sections of Vijaya et al. (2012)
the terms “ovo-spermathecal duct” and “ovarian vesicle”.
Spermathecae: As just noted under ‘Female organs’, there is an atrium with muscular
and enclosed in a sheath; at their junction a long oviduct attaches leading to the ovisac
which is opposed by a small saccular outgrowth. The whole or just part of the structure
may be referred to as a ‘fertilisation chamber’ as it functions for internal fertilisation of
eggs with sperm, presumably before transfer of the embryos to the cocoon.
Prostates: Large pair of digitiform euprostates, with white muscular sheen from 18
extending to 23; acutely muscular enlargements of loop of paired sperm ducts which
attach to apex of copulatory chamber mound centrally. As noted, a smaller blind duct
— the Y-shaped gland — attaches to the base of the mound mesially, although Beddard
Other internal features: Small saccular ‘brown bodies’ formed from coelomocytes
were observed loose in coelomic cavities from 7 posteriorly; these may enclose shed
Pontoscolex; the function
in both cases is unknown.
The gut contains soil and/or organic matter (depending on habitat) — this species
appears to be an adaptive feeder and will survive in unaltered soil (as noted) but also
Cocoons: Dark coloured with adhesions, tapered lemon-shape with one side usually being
incubation and hatching data); may contain from one to eight hatchlings (Gates 1982).
Distribution (Fig. 3): After Michaelsen (1903: 122); Gates (1942: 98, 1972: 52, 1982: 72):
West African origin from Upper Guinea plain or coastal forest including Sierra Leone,
BLAKEMORE: EUDRILUS EUGENIAE 533
Liberia, Ivory Coast, Ghana, Togoland (Benin), Nigeria, Cameroon, Gabon and the
Congo; transported and peregrine to many tropical countries such as Madagascar and
the Comoros Islands (e.g. Anjouan), Seychelles (Gerlach 2011), Sri Lanka and India
(Michaelsen 1903; Stephenson 1923: 486; Dhiman & Battish 2005), and New Caledonia;
the Americas: [e.g. Gates (1982: 74) said it owes its North American distribution since
lower 48 United States, such as Florida, Alabama, Georgia, Texas, and even to Hawaii,
(Rodriguez-Aragones 1999), Suriname (Horst 1887), Panama [from 1896 — Eisen
(1900: 135) said: “Judging from the number of specimens in the collection, this species
as an introduction from the then ‘British Honduras’ noted by Gates 1982), Venezuela
(e.g. roseus), Guyana, Colombia (Feijoo et al. 2004), Paraguay (Schuldt 2009), Brazil;
Virgin Islands (Michaelsen 1903, 1910; Gates 1942, 1972), Cuba (Gates 1972; Alvarez
& Rodriguez-Aragones 2010), Bahamas, Antilles (Gates 1942: 99) and Guadeloupe
(Csuzdi & Pavlicek 2009 — who found it in a natural setting indicating it may have
become feral there as it is on St Helena); also the Atlantic: Bermuda (as E. erudiens),
St Helena (type-locality by introduction), Cape Verde (from where it was introduced
1903, 1910). Elsewhere in America, Gates (1982: 72–74) explained in some detail how
“can of worms (bait) inadvertently left behind” and cultured by the camp owner (a Mr
Fig. 3. Distribution map from Michaelsen (1903: chart 1) (hash marks family distribution). Note that New
Zealand was in error but other records outside its West African homeland are due mainly to human
transportation and the worm’s acclimatisation; early Caribbean and Latin American introductions
possibly relate to the 16th – 19th century Atlantic slave trade.
534 AFRICAN INVERTEBRATES, VOL. 56 (3), 2015
T. Baker), eventually shipped to all of the USA and Canada where it has been cultured
both indoors and outdoors.
in 1991 (Blakemore 1994, 1999) with stock (surface sterilised cocoons) originally
In Europe it was introduced to Hamburg with plants from the West Indies (Michaelsen
1903: 12) and to Kew Gardens in Wardian cases from British Guiana (Beddard 1906). It
is rarely reported from northern European glasshouses by Sims and Gerard (1985), albeit
rarely, e.g. from Denmark (Blakemore 2007) and eastern Europe, Hungary (Csuzdi et al.
2007 ); also maintained in laboratory cultures, e.g. Vigo, Spain (Dominguez et al. 2001).
Plisko (2010) notes that it was deliberately introduced to South Africa (RSA) by
Reinecke and Viljoen (1988) from Germany in stock originating in West Africa and
that this species is now widely used in RSA farms and is “adapting well to habitats in
this country” suggesting its naturalisation there.
Eudrilus eugeniae is stated to be newly introduced to Egypt (Medany & Yahia 2011:
20), but what this paper actually says is: “Four types of earthworms were brought
to Egypt from Australia: Lumbriscus Rubellus (Red Worm), Eisenia Fetida (Tiger
Worm), Perionyx Excavatus (Indian Blue), and Eudrilus Eugeniae (African Night
Crawler)”. However, Lumbricus rubellus Hoffmeister, 1843 has never been proven a
vermicomposting worm (Blakemore 1999, 2002), thus it is likely only three species or
fewer were involved. Eudrilus
At least one worm farmer in Valparaiso, Chile and a technician (Mr Reinaldo Plasen-
cia) in Nicaragua claim to rear Eudrilus (“la lombriz africana”) sometimes misspelled
“Fudrillus spp” (Lumbricultura 2014; Monographias 2014), which would both be new
national reports. Mr Enzo Bollo Tapia (pers. comm. 2014) communicated that it can be
cultivated in Ecuador, Colombia and Peru but that Chile is unsuitable for its survival
due to climate, although he did experiment there.
Introduced to the Philippines for vermicomposting in the 1980s, E. eugeniae is now
distributed in worm-beds on farms over the whole country. A report of its spreading to
some mountainous inland areas via agro-forest strips of Negros Occidental by Flores
(2007) is unsubstantiated as there is no proof that E. eugeniae itself was found. The
report just says “Eudrilus” based on a novice’s key to families. It is also newly reported
from Thailand, from an unpublished DNA barcode submission to GenBank in 2010/2011
(see Appendix) and recent reports from there (e.g. Malliga 2010; Loongyaii et al. 2011).
Eudrilus is used for soy bean residues and rice husks vermicomposting in Malaysia
(e.g. Lim et al. 2011; Shak et al. 2014) with the worms apparently imported as cocoons
from India. It is also reported from Indonesia where vermiculture operations in Solo,
Central Java are advertised (e.g. Indonetwork 2014; Cacinglumbricus 2014). This has
West Java (Andy Darmawan pers. comm. via email Nov. 2014). Recent reports from
Vietnam are from the provinces of Lang Son and Cao Bang by the Research Institute for
et al. (2011); AFSPAN (2012) but mispelt “Eudrilus euganaie
New Zealand records by Beddard (1895: 149), repeated by Michaelsen (1900,
1903), Hutton (1904: 355), Gates (1972), and Sims and Easton (1985) were stated by
Thompson (1922: 359), Benham (1950) and Lee (1959: 365) to be an error introduced
BLAKEMORE: EUDRILUS EUGENIAE 535
when Beddard (1891, 1895: 149) somehow mistook for Eudrilus eugeniae Smith’s
1886 report of Endrilus [sic lapsus for Eudriluslevis [= Octochaetus? levis (Hutton,
Zealand also failed to locate this species there (e.g. Blakemore 2012a).
The claim from the French islands off the coast of Newfoundland (St Pierre and
E. lacazii by Perrier (1872) was disputed by Gates (1982: 72), although
Philippines and South Africa), from a few southeast Asian countries neighboring
Vietnam, or yet from China/Taiwan.
Locality: Specimens were collected from worm farms in Brisbane (1991) and samples
sent to the author from Mackay, Qld (1992), and Menai, NSW (1996) [now in CSIRO/
but only close to worm beds; neither was it located ferally in surveys on Negros Island
(pers. obs. 2009– 2014, cf. Flores 2007).
Habitat: Originating in shaded savannahs of West Africa, it now thrives in worm beds
on worm farms; it is reported in natural high moisture/organic sites such as waterfalls
Miagao, Philippines (pers. obs. Feb. 2014).
Behaviour: Hatchlings are reported to sometimes return to the cocoon when alarmed.
Active with a rapid escape response when disturbed, but if captured the adult worms
become placid and can be readily handled. The species will wander at night, leaving
that are shaped similar to a scorpion’s stinger (see Fig. 2).
The genus Eudrilus is bi-parental, being characterised by internal fertilisation preced-
ing cocoon production (Sims 1964, 1987). Initial life studies are relatively recent, for
example by Neuhauser et al. (1979), who found the best growth on horse manure or acti-
vated sewage sludge. Its life span can be 1–3 years, with Eudrilus eugeniae possessing
a life cycle that ranges from 50– 70 days, with sexual maturity reached at 35–50 days
in culture (Viljoen & Reineke 1989). Reineke and Viljoen (1988) reported that in a
cattle manure substrate at 25°C, cocoons produced by adult worms between the ages of
70– 100 days were incubated for ca. 17 days before producing a mean of 2.7 hatchlings
per cocoon (range 1–8) with 84 % hatchling success. Dominguez et al. (2001) had similar
3.6 cocoons per week with 2.2 viable hatchlings per cocoon (= 6.5 hatchlings per worm
clitella at between 35– 45 days; worms with fully developed clitella copulated readily
536 AFRICAN INVERTEBRATES, VOL. 56 (3), 2015
and the formation of cocoons started within 24 hours after copulation, continuing for
up to 300 days. In India, Nagavallemma et al. (2004) recorded an 18-fold increase in
population (from 55 specimens to 1,007) on legume leaf/cow dung substrate in three
months, the highest of three composting species they tested. Also in India, Vasanthi et
mixed with sawdust and cattle manure at 26°C. These rates of growth and reproduction
are amongst the highest currently reported for any earthworm.
Parasites and disease resistance
Gates (1982: 74) states that, unlike in most earthworm species, parasitic protozoans
had not been reported. Internal nematodes (parasitic, paratenic or commensal) are known
(e.g. Gates 1974: 74; Poinar 1978; del Valle & Rodriguez 1988; McNeill & Anderson
1990; Spiridonov 1992), but this species is supposedly disease-free apart from records
of ammonia lesions on the clitellum (Gerasimov 2007). The functions of the ancillary
glands of the female and male organs are not fully worked out; possibly they produce
nutrients for eggs/sperm or are in part copulation sentinels preventing intromission of
parasites or other disease vectors. Neither is the function of the supra-intestinal glands
understood, as already noted.
Gates (1982: 74) reports ‘head’ regeneration as well as more easily observed posterior
et al. (2014), thus it
is plausible for this species to get two viable worms from a single ‘individual’ as with
some other species reported by Blakemore (2001).
Ecology and economics
Ecology of the three most common aerobic composting worm species, Eisenia fetida
(Savigny, 1826), Eudrilus eugeniae and Perionyx excavatus Perrier, 1872, that are most
often bred in worm farms and fed on household vegetable wastes or animal manures, are
detailed in reports by Graff (1982), Sabine (1983), Kale and Bano (1991), Reinecke et
al. (1992), Kale and Sunitha (1993, mispelt “Sunita” in Edwards 2004 : 388, table 19.3)
and by Dominguez et al. (2001). Comparative studies generally show E. eugeniae to be
a most productive species in tropical zones or under cover in temperate regions (where
Eisenia fetida, as noted below. For example, Gates (1982: 74) says it is the preferred food
for duck-billed platypuses [Ornithorhynchus anatinus
In Cuba, India and the Philippines, this worm is favoured most for producing vermi-
compost fertiliser for organic farming, whereas in North America and Australia the
‘African Nightcrawler’ or ‘ANC’. The current studies noted a propensity to escape from
potential for mobility, there were no records of natural colonisation for North America
(Gates 1958, 1972, 1982) or Australia, and such records from New Zealand are now
BLAKEMORE: EUDRILUS EUGENIAE 537
known to be mistaken identities. As Gates (1958: 10) said: “This species, originating in
tropical Africa and until very recently known only from the tropics, has been raised and
distributed in the United States for several years by earthworm culturists. Sales appear to
be mostly to anglers for bait. Escapes of live specimens into natural environments must
have been numerous. As yet, however, there are no records to indicate acclimatization
and permanent colonization in mainland states.” This was slightly counter-indicated
by his later record of specimens from soil under oak trees at Vero Beach Laboratories,
Florida (Gates 1982: 72).
Nevertheless, the spread of Eudrilus eugeniae
is a widely cultivated species in Brazil too and there are some records of its survival
away from worm farms, mainly in areas of high organic matter, but also in gardens
& Pavlicek 2009). A novice report from mountainous forests in Negros Occidental,
Philippines (Flores 2007) is unsubstantiated, being based on a simplistic, non-specialist
key only to families, and re-surveys (unpub.) by the current author have not found it
far from culture beds.
Preferring bedding material rich in organic matter in culture, this savannah worm
also survives in unamended soils (M’ba 1978); and Blakemore (1994) successfully
maintained it for six months with reproduction in mesocosms of heat-sterilised but
unamended clays (vertisol and kraznozem) and sandy (podzol) soils in the glasshouse.
Parthasarathi (2007) showed that over a year it will grow in clay loam but not as well as
in composts where its biomass may be six times as high; thus it is considered adaptable
to a wide range of soil types, unlike most other highly restricted earthworms that co-
evolve with their soils (Michaelsen 1922).
Such environmental tolerance was investigated by way of soil selections by Madge
(1969) who introduced Eudrilus eugeniae and another tropical species to gradients
of soil texture and found a marked preference for the 0.25 mm particle size fraction
Blakemore (1994) showed that it, along with Eisenia fetida, had a tendency to select
soil amended with manure when given a choice and compared to other species; it was
found in clay soil rather than loam or sandy soil in 25 % of all its observations. Habitat
E. eugeniae and another
tropical African species to a soil moisture gradient and found 65 % of the earthworms in
the 12– 17 % moisture sectors after 48 hours; tolerated pH range was between 5.6– 9.2
and for temperatures, he found an optimal range between 23°C and 31.5°C. In growth
experiments, Viljoen and Reinecke (1992) reported that no E. eugeniae juveniles survived
below 12°C or above 30°C, and optimal temperature for growth and reproduction was
around 25°C. Attempts to establish it in natural environments show that the worms do
well until the temperature drops to 40°F (4.4°C), at which time they die (Gates 1972:
52). Sims and Gerard (1985, 1999) say that temperatures below 10°C are not tolerated
and that the optimum breeding temperature range is 21–27°C. Domingues et al. (2001)
80– 82 % and 25–30°C. These moisture and temperature levels correspond well to
a light sandy loam (10 % clay) and the preferred temperature was 25°C, in gradient
cylinders laid horizontally to circumvent depth affects.
538 AFRICAN INVERTEBRATES, VOL. 56 (3), 2015
Nature of Eudrilus casts
Eudrilus eugeniae is plentiful in the coastal, shaded savannah grasslands of its West
African homeland where copious surface pellets are produced — remarkably, up to 140 t ha-1
-1 per annum according
to Gates (1972: 52), during the rainy season only.
In a series of mesocosm soil cores trials with combinations of species, soils and
test crops, Blakemore (1994, 2008a) determined that E. eugeniae produces distinctive
container edges where the worms burrowed, and had the highest surface cast production
for a high casting rate of E. eugeniae estimated by Cook et al. (1980) of up to 2.43 g soil
per gram fresh weight worm per day. The casting rates in the 23 cm diameter (0.0434 m2)
by 33 cm deep mesocosms were as high as 610.5 g per pot in kraznozem soil in just six
the rate reported by Madge (1969).
Blakemore (1994, 2008a) observed that, where casts for this species were deposited
on the surface, adventitious plant roots sought them out and that casts fallen over the
base too; when the roots were lifted cast pellets looked like miniature bunches of grapes
dangling on a vine. This may be explained by Eudrilus eugeniae
higher nutrients and trace elements in its casts compared to the clay soil matrix, especially
nitrate-N, K and Zn, which were about twice that of the soil (Blakemore 1994, also
Table 1), and possibly other plant attractant compounds.
Effects on plant yield and soil moisture after deliberate introduction of this species
cores ranging from +21–74 % for oats and +15–50 % for grass shoots (and up to +50 %
by surface casts and also drainage in worm burrows. As one of a dozen candidate species
pasture grass yield by 83.9 % (i.e. nearly doubled compared to controls). However,
these preliminary results were considered inconclusive due to background variation and
lack of survivors after one year during a particularly severe drought in tropical eastern
Australia that precluded irrigation even from cattle stock reservoirs.
Inadvisability of deliberate introductions of alien species
The ethics of releasing an exotic species such as Eudrilus eugeniae into the Australian
this was an objective of the project that aimed to enhance improved pasture production
naturally (Blakemore 1994). Possible risks were considered acceptable on the grounds
that this species was present in Australia, having been legally imported from Canada by
bait, including in the Mundubbera township nearest to the release site (pers. obs.). This
than being parthenogenetic) and the upland release area, classed as dry, arid sub-tropical,
BLAKEMORE: EUDRILUS EUGENIAE 539
was several kilometres from a water course that may have aided dispersal. Thus the
likelihood that the species would persist due to the single release event, especially
in the prevailing drought, was considered negligible. However, increasing concerns
about the spread of alien species would mean it is inadvisable to contemplate such a
release or deliberate introduction in the future in Australia or elsewhere. Commercial
solutions for the worm bait/compost market could be found from the pool of native
species, some of which are already bred as bait in Australia (Blakemore 1999, 2012b)
but, again, redistribution of these natives is also ill-advised albeit less objectionable and
Sruthy et al. (2013) determined that the intestinal microbial populations of Eudrilus
eugeniae in the foregut, mid and hindgut were dominated by bacteria, actinomycetes
and fungi, respectively. These authors reviewed the diversity of types and number
of these microbes, plus yeasts and protozoans in the casts of E. eugeniae and other
vermicomposting species reported on by other researchers (see next section below).
Use of earthworms in vermistabilisation of sewage sludge and other wastes have had
Composition of Eudrilus eugeniae casts produced from two types of unamended soils: (A) a sandy podzol
and (B) a clay vertisol (Blakemore, 1994: tables 4.3.22/23) compared to (C) 100 % ANC vermicast
source soil medium; 1 % = 10,000 mg/kg or 10,000 ppm (parts per million); ‘~’ = conversion estimates.
Component (A) Sand soil casts (B) Clay soil casts (C) 100% Vermicast/
Moisture - - 30 %
Organic matter 36 %
C:N Ratio 15:01
Total C ~21–28 %
Total N* 1.89 %
P2O5 (from P-Bicarb)* 2.49 %
K2O (from K total)* 1.40 %
Ca 5.09 %
Fe - - 2.63 %
Mg 0.17 %
MEAN ratio to soil -
*Total macronutrients N-P-K are generally less important for composts that are microbially activated.
540 AFRICAN INVERTEBRATES, VOL. 56 (3), 2015
favourable results when attempted in various regions (e.g. Neuhauser et al. 1988;
Blakemore 2000b, c). Monroy et al. (2008) showed that processing of pig manure slurry
with E. eugeniae eliminated nematodes and reduced coliform bacteria by up to 98 %.
Composition of ANC vermicompost
The chemical and microbial characteristics of E. eugeniae vermicast/vermicompost
differ depending upon the nature of the substrate and the age of the casts. Chemical
composition is determined by the source material on which the worms are fed but is often
enhanced in terms of natural plant nutrients, these changes relating to physical, chemical
and microbial activities during and after passage through the worms’ intestines. Table 1
summarises data from experiments by Blakemore (1994: tables 4.3.22, 4.3.23) that
found that plant nutrients in casts of clay soil increased by about +20 %, whereas casts
from a sandy soil were depleted in nutrients (by -20 %) probably due to assimilation
by the worms, although structural characteristics of the soil matrix were improved by
the worm activities too, increasing plant yields as noted above. Several authors, have
et al. (1988). For
instance, Nagavallemma et al. (2004) found generic vermicomposts to have higher
percentages (nearly double) of both macro- and micronutrients and higher microbial
activity compared with garden composts, plus they detected these plant-growth-
promoting agents. Data from Parthasarathi (2006) is summarised in Table 2.
to mineralisation and the gradual release of nutrients, as well as plant-growth-promoting
enzymatic agents. As vermicompost ages, this biological activity declines, but the
vitamin content may double with time (Prabha et al. 2007). Studies by Parthasarathi
and Ranganathan (2000) showed that enzymes (cellulase, amylase, invertase, protease
and phosphatase) declined as the casts aged. Parthasarathi (2006) found that Eudrilus
p<0.05) in microbial population (fungi + bacteria + actinomycetes) and dehydrogenase
enzyme activity in fresh casts, leading to enhanced nutrient mineralisation, but this
activity gradually decreased over the period of a month as the casts aged (see Table 2).
Effects of ANC vermicompost on plant yield
A study by Kale et al. (1992) commented on applications of Eudrilus eugeniae ver-
-1 plus half of the recommended
are similar to those reported by Pontillas et al. (2009) from the Philippines. However, the
Blakemore (2000a) which found that a partly organic section of Haughley Farm was not as
productive as a wholly organic section. Unpublished data from Kahariam organic farm in
the Philippines show that ANC vermicast application to paddy at rates of just 1.5–3 t ha-1
-1, well above the typical local
range of 20– 90 cavans ha-1 but without the need for chemical additions (Mr Danny
applying up to 30 t ha-1 Eudrilus vermicompost yields 90 t ha-1 (Mr R. Peñalosa, pers.
comm.), above the regional average yield of 50 t ha-1, i.e. +80 %. In current studies by
BLAKEMORE: EUDRILUS EUGENIAE 541
the author, the resident earthworms on both farms were enhanced in terms of biomass
and biodiversity when compared with neighbouring farms that use conventional chemical
using vermicomposts as primary fertilisers.
Composition of sh bait, stock feed and worm meal
Worms can be fed to stock directly, or dried and added to worm meal. Composition
is provided by several authors, for example Hertrampf and Piedad-Pascual (2000) who
show 85.3 % moisture in E. eugeniae worms with 56 % protein when dried, and ten
essential amino acids plus macro and trace minerals of freeze-dried vermimeal. Although
eugenia are not presently available.
Radioactivity and biocide effects
Eno (1966) found E. eugeniae to be less susceptible than Lumbricus terrestris Linnaeus,
1828 to irradiation in the range 16– 64 kR. In Nigerian soils, this worm had much higher
concentrations of DDT and its products, compared with the surrounding soil, and its
production of surface casts virtually ceased in DDT-treated plots, which was considered a
contributory factor to the overall decline in fertility of these plots (Cook et al. 1980). The
concentration of heavy metals (Cu, Zn, Pb, Cd and Hg) in the tissue of E. eugeniae fed
on municipal wastes accumulated beyond acceptable levels for protein-meal production
(Graff 1982), although this trait could be utilised for soil bioremediation.
Pharmaceutical or cosmetics uses
coagulent or thrombolytic drugs to treat and prevent cardiac and cerebrovascular diseases
in heart and stroke patients. In original 1982 Japanese patents (e.g. US4568545A) extracts
were said to come from “Lumbricus rebellus” (sic), but Indian researchers (Sharma et al.
eugeniae as well as Eisenia fetida
anticoagulant properties of E. eugeniae and its extracts have been variously studied (e.g.
Mathur et al. 2010a; Packia Lekshmi et al. 2014). When Eudrilus eugeniae was dried
and powdered, it produced antimicrobial responses of between 45– 90 % inhibition on
agar plates when assayed against seven human pathogens (Anjana et al. 2013). Extracts
from this worm showed antibacterial and antifungal properties that varied depending
on the formulation (Mathur et al. 2010b). Preliminary studies by Shobha and Kale
(2008) found indications of possible control of plant soil-borne fungal and bacterial
pathogens using E. eugeniae et
al. (2013a) using a paste from this worm to inhibit the growth of resistant/recalcitrant
human pathogens of bacteria such as Staphylococcus aureus Rosenbach, 1884 (‘Golden
Microbes in ANC vermicompost (after Parthasarathi, 2006: table 1); CFU = Colony Forming Units.
Casts Bacteria (CFU g-1) Fungi (CFU g-1) Actinomycetes (CFU g-1)
Fresh casts 444
30-day-old casts 444
542 AFRICAN INVERTEBRATES, VOL. 56 (3), 2015
staph’) and of fungi such as Candida albicans (Robin, 1853) (‘Thrush’), indicating its
Emulating the works of Cooper et al. (2004a, bE. eugeniae
as the source, Dinesh et al. on
cancer cell lines of human HeLa cells, colon cancer cells, WBC malignant tumour cells
and brain tumour cells, with reduction by up to 33 %. Such studies indicate novel use
of this earthworm for treatment of cancers.
Azmi et al. (2014) recently found earthworm extracts, including those from E.
eugeniae, to have anti-wrinkling properties as potential new ‘anti-aging’ agents.
Eudrilus eugeniae is a remarkably versatile vermicomposting species of the tropics
or indoors in temperate regions. It is useful for recycling soil organic matter (i.e.
stock-feed protein, bioprospecting for pharmaceuticals/cosmetics or perhaps silk produc-
bait. Its geographical range and the applications of its products are rapidly expanding,
with a summary of its vermiculture potential provided by Li et al.
similar Eudrilus pallidus (Michaelsen 1892: 216) from Accra, Ghana [?syns. E. buettneri
Michaelsen, 1892; Eudrilus ifensis
species E. pallidus atakpamensis Michaelsen, 1913 and E. simplex Michaelsen, 1913
unlike the 7,000 other earthworm species currently described but mostly neglected, and
these perhaps just a fraction of the total numbers in nature (Blakemore 2012b). Thus,
of earthworm ecology still remains largely open and unexplored.
Preliminary taxonomic work was undertaken during PhD studies at CSIRO Tropical Agriculture, Brisbane
under an Australian RIRDC grant from 1991–1993. Inspection of Natural History Museum, London
NIBR under the auspices of Dr Wonchoel Lee of Hanyang University, Seoul whence Dr Seunghan Lee
in Batangas, Philippines. Comments of editors and anonymous referees helped improve this contribution.
BLAKEMORE: EUDRILUS EUGENIAE 543
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