Review of the balantine genus Udeina Michaelsen, 1910 with descriptions of six new species in South Africa (Oligochaeta : Acanthodrilidae, Acanthodrilinae)

2004·1.3 MB·English

by  Plisko, Jadwiga Danuta

African Invertebrates Vol. 45 Pages 287–313 Pietermaritzburg December, 2004
Review of the balantine genus Udeina Michaelsen, 1910 with
descriptions of six new species in South Africa (Oligochaeta:
Acanthodrilidae, Acanthodrilinae)
by
Jadwiga Danuta Plisko
(Natal Museum, P. Bag 9070, Pietermaritzburg 3200, South Africa and School of
Botany & Zoology, University of Natal, P. Bag X01, Scottsville 3209, South Africa;
[email protected] or [email protected])
ABSTRACT
A brief historical overview of the South African acanthodriline earthworms is given. The taxonomic
position of the genus Udeina Michaelsen, 1910 is reviewed. An inclusion of the Udeina species, characterised
by balantine arrangement of male reproductive organs in the genus Parachilota Pickford, 1937 as proposed
by Ljungström (1969, 1972), is not accepted. Udeina is retained and redefined, and eleven species are now
included. The indecisive status in the literature of two species, U. kinbergi (Michaelsen, 1899) and
U. reichei (Ude, 1905), is discussed. New material of U. reichei has been studied, and the validity of both
species confirmed. The type material of U. montanus Pickford, 1937, U. pliskoae Zicsi, 1998, and
U. stuckenbergi Zicsi, 1998 was examined, and new observations, comments, and illustrations are provided.
Six new species—U. adriani, U. anneae, U. mapelane, U. nkandla, U. petrosi and U. qudeni—are described
and illustrated.
Identification keys are provided to the currently recognised genera of South African Acanthodrilinae and
to species of Udeina.
The species name U. stuckenbergeri erroneously published by Zicsi (1998) is corrected to U. stuckenbergi.
INTRODUCTION
Although Pickford (1937), Ljungström (1969, 1972), Zicsi & Pajor (1992), and Zicsi
(1998) have supplied considerable data on acanthodriline diversity in this country, the
indigenous South African species still require substantial study. The known
Acanthodrilinae are probably only a minor part of those that may be discovered. The
present paper forms a part of a survey of acanthodrilines in South Africa, based on the
collection in the Natal Museum.
For those Acanthodrilinae monographed by Pickford (1937), features of the male
reproductive organs were used to separate genera: 1/ the primitive, acanthodriline
arrangement of paired prostatic and male pores; 2/ microscolecine and balantine
arrangements manifesting reductions in the number of prostates, and relocation of
prostatic and male pores. The morphological nephridial differentiations, described as
vesiculate or avesiculate, have been recognised as supportive diagnostic characters
separating vesiculate microscolecine species from avesiculate balantine species. For
all the acanthodrilines recorded from South Africa, the majority being indigenous
species, five genera have been proposed: Eodrilus Michaelsen, 1907, Chilota
Michaelsen, 1899b, Microscolex Rosa, 1887, Parachilota Pickford, 1937, and Udeina
Michaelsen, 1910. Gates (1959) and Ljungström (1969, 1972) disagreed with the
separation of the proandric, avesiculate, balantine species from proandric, avesiculate
acanthodriline Parachilota species, suggesting that the balantine reductions might be
only variations occurring in those Parachilota species characterised by more primitive
287
288 AFRICAN INVERTEBRATES, VOL 45, 2004
acanthodriline male reproductive organ arrangements. Zicsi (1998) did not follow their
suggestion, recognising the genus Udeina for the avesiculate species with the balantine
prostatic reduction, known only from South Africa. This supported his interpretation
that the genus Yagansia, Michaelsen, 1899 is accepted for those South American
vesiculate species with microscolecine prostatic reduction. In the present paper, the
validity of Udeina is recognised for South African species, and supported by additional
evaluation of its generic features, notably the proandric condition associated with
reductions to one pair of spermathecae and one pair of prostates, and constant occurrence
of these characters.
The taxonomy proposed by Pickford (1937) is accepted here, although following
Zicsi’s (1998) recommendation, the South African species assigned by her to Eodrilus
are accepted in Eodriloides Zicsi, 1998, supported by similar exclusion of Australian
species, including the type-species Eodrilus cornigrave Michaelsen, 1907, from Western
Australia, by Jamieson and Dyne (1976). The present paper comprises new taxonomic
observations, evaluation of characteristics of the genus Udeina, and descriptions of six
new species.
MATERIAL AND METHODS
During my extended survey of earthworms in South Africa, a collection of
acanthodrilines was assembled, and specimens with balantine reductions of the anterior
or posterior pair of prostates were noted. An additional substantial material of
acanthodrilines donated to the Natal Museum by Dr A. J. Armstrong of the Biodiversity
Division, Ezemvelo KZN Wildlife, and by the Mammal Research Institute, University
of Pretoria, supplied more individuals with these characters. This stimulated my attention
to evaluate South African proandric species, characterised by only one pair of
spermathecae, balantine reduction of one pair of prostatic glands, and relocation of
male pores.
The type material of known species Udeina montanus Pickford, 1937, U. stuckenbergi
Zicsi, 1998, and U. pliskoae Zicsi, 1998, was included. No type material of U. kinbergi
(Michaelsen, 1899) and U. reichei (Ude, 1905) has been examined. The types of both
species were restudied by Michaelsen (1907, 1910, 1912) and Pickford (1937). Both
authors confirmed that although the material was in poor condition, balantine reductions
were evident. However, the exact locations of prostatic and male pores in the types
remain uncertain, as neither author could definitely locate them, and probably further
examination could be uninformative due to decomposition of the specimens.
A small sample containing two slightly decomposed adults and seven juveniles,
collected on a bank of the Vaal River, possibly near the type locality of U. reichei, and
identified by Ljungström (1972) as U. kinbergi, was examined by me and found to be
U. reichei.
The types of new species described below—U. adriani, U. anneae, U. mapelane,
U. nkandla, U. petrosi, and U. qudeni—and of already described U. pliskoae and
U. stuckenbergi, are preserved in 75% ethanol and housed in the Natal Museum.
U. montanus is in the South African Museum, Cape Town, South Africa. The
descriptions of all species are based on type specimens, and comparisons with available
additional material are made. Description of colour is based on specimens which may
have lost intensity during preservation and through storage in alcohol or formalin. The
PLISKO: UDEINA IN SOUTH AFRICA 289
number of body segments was counted from the anterior segment in which the
prostomium is located, and presented in Arabic numerals. Intersegments and septa
location are designated by a slash (e.g. 7/8). Dorsal longitudinal dissections were
performed for study of the internal anatomy, and the photographs were taken under a
Wild stereo microscope Type 181300. Penial setae were observed in glycerol jelly,
permanently mounted in Berlese fluid, and photographed using photo-camera Wild
MPS 51 attached to Wild Heerbrugg microscope.
Locality data and biological notes were taken from labels; information on the biotopes
was often extended by personal comments from Dr A. J. Armstrong, or taken from
Pooley and Player (1995). The species U. adriani, U. mapelane, U. nkandla, U. petrosi,
U. qudeni and U. stuckenbergi are known from relatively undisturbed locations,
suggesting that South African acanthodriline earthworms prefer pristine, natural biotopes,
indigenous grasslands and forests. Probably more species will be found in other protected
areas of South Africa.
Generic characters of South African Acanthodrilinae reported by Pickford (1937) are
presented in Table 1. Keys are provided to recognised genera of South African
Acanthodrilinae and to the species of Udeina.
Glossary, abbreviations and definitions of terms used in the text:
1/n immediately followed by a figure refers to that part of segment covered by clitellar
tissue.
TABLE 1
Generic characters of South African acanthodriline earthworms
presented by Pickford (1937).
Genus Microscolex Eodrilus= Chilota Parachilota Udeina
Eodriloides
Spermiductal/ holandric, or holandric proandric proandric proandric
testes funnels proandric with
rudimentary
funnels in 11
Spermathecae one pair, or two two pairs two pairs two pairs one pair
pairs, or absent
Spermathecal 7/8 and 8/9; 7/8 and 8/9 7/8 and 8/9 7/8 and 8/9, or 7/8
pores or 7/8 or 8/9; on the anterior
or absent part of 8 and 9
Male pores 17 or 18 or 18 or 17/18 18 18 or 19 17 or 18
absent
Prostates one pair, or two two pairs two pairs two pairs one pair
pairs, or absent
Prostatic 17 and 19 17 and 19 17 and 19 17 and 19, 18 or 19
pores or 18 and 20
Male repro microscolecine acanthodriline acanthodriline acanthodriline balantine
ductive organs
Excretory vesiculate avesiculate vesiculate avesiculate avesiculate
system
Indigenous to one species 15 species 13 species 52 species two species
SA
Introduced to two species none none none none
SA
290 AFRICAN INVERTEBRATES, VOL 45, 2004
Acanthodriline arrangement of the male reproductive organs, or shortened as one word:
acanthodriline – refers to location of two pairs of prostatic pores of the tubular
prostates in segments 17 and 19 respectively, and male pores present in segment
18. This condition is usually associated with two pairs of spermathecae, and
spermathecal pores occurring in intersegmental furrows 7/8 and 8/9 respectively.
This arrangement is considered as primitive in the Acanthodrilidae.
Avesiculate – referring to a holoic single nephridium without dilation. The term was
used by Pickford (1937) to describe the nephridia observed in acanthodriline species
allocated to Eodriloides, Parachilota and Udeina.
Balantine – originally the term referred to a reduction of anterior pair of prostates and
posterior pair of spermathecae, occurring in species characterised by acanthodriline
condition; one pair of prostates occurs in segment 19, and a pair of spermathecae
of segment 8 has its pores located in intersegmental furrow 7/8. The present
understanding employed in this paper, refers to acanthodriline, avesiculate species,
with one pair of prostates, with prostatic pores in 17 or 18 or 19, and with one pair
of spermathecae with spermathecal pores in intersegmental furrow 7/8 or 8/9. Male
pores may occur in 16 or 17 or 17/18 or 18.
Clitellum – the region of the body-wall formed from glandular cells, which secrete
material to form a cocoon. It may be saddle-shaped or ring-shaped, although the
shape may depend on the phase of sexual maturity. The position of clitellum in
most South African Acanthodrilinae species is usually constant, but intraspecific
variation occurs in some species.
Figures followed by a stroke (for example 7/8) indicate a position between segments or
septa.
Holandric – refers to number and position of testes and testes funnels; indicates presence
of two pairs of testes and two spermiductal funnels occurring in segments 10 and
11 respectively; (compare proandric).
Holoic – (old terms: holonephric or meganephric); refers to the paired excretory organs,
as simple nephridia in each segment of the body except the pharyngeal segments.
Holoic nephridia are observed in all acanthodrilids. Pickford (1937) noted in some
species that the single nephridium may possess variable dilation forming terminal
bladders, and on this basis, for the species recorded in South Africa, two groups
were established: the avesiculate nephridia found in Eodriloides, Parachilota and
Udeina, and the vesiculate nephridia in Chilota and Microscolex.
JDP – J. D. Plisko.
KZN – KwaZulu-Natal province of South Africa.
Male pores – ectal ends of vasa deferentia. In majority of the South African
acanthodrilines male pores are situated laterally to the b setae; only in two known
species (P. vanhoffeni and P. bavenda) approximating towards the mid-ventral line.
In some species may be fused with prostatic pores.
Microscolecine – originally this term referred to a reduction of posterior pair of prostates
and reduction of anterior pair of spermathecae, observed in the species with
acanthodriline male reproductive organs. In the vesiculate species recorded in South
Africa, the microscolecine characters have been observed only in two introduced,
and in one endemic species. Variability in reduction and relocation of male organs
were noted.
PLISKO: UDEINA IN SOUTH AFRICA 291
NMSA – Natal Museum, Pietermaritzburg, South Africa.
Penial seta (pl. penial setae) – specialised seta associated with prostatic pore; development
of penial seta depends on maturation of the individual; after the sexual phase, the well
developed seta is discharged, being gradually replaced by a new precursory seta, which
is progressively modified. During seta formation, precursory setae come to resemble
the adult penial setae except in size, and their shapes may be observed. In the majority
of known South African acanthodrilines, the penial setae were observed and often
described; however, the state of their maturity was not always indicated. The taxonomic
importance of penial setae is considerable.
Penial setal retractor muscles (pl. retractors) – the muscles originating from the body
wall dorso-laterally, usually behind the penial seta and activating them; maturity
of retractors depends on maturation of individuals. The site of origin can be of
taxonomic value.
Proandric – refers to number and position of testes and their funnels; indicates the
presence of a single pair of testes and spermiductal funnels in segment 10; (compare
holandric).
Prostate (pl. prostates) – paired glands associated with male reproductive organs; each
consists of a prostatic gland and muscular prostatic duct. In most Acanthodrilidae
the prostatic gland is tubular; according to Pickford (1937), it originates as an
ectodermal invagination lateral to seta b; the number and position correspond to
the number and position of prostatic pores, and for acanthodrilids it is an important
taxonomic character. In South African species always tubular.
Prostatic duct – extended part of prostatic gland, usually muscular; its ectal part opens
into prostatic pore.
Prostatic gland – in South African Acanthodrilinae, the glandular, tubular part of prostate;
may be confined to one segment or extended through several segments.
Prostatic pore (pl. prostatic pores) – ectal part of prostatic duct. In the endemic South
African acanthodrilines the pores are situated lateral to the b setae; only in two
species (P. vanhoffeni and P. bavenda) the prostatic pores approach the mid-ventral
line. South African indigenous mature specimens usually have the pores connected
with the male pores by the seminal grooves.
Salivary glands – pharyngeal structures; in South African acanthodrilines usually
extending backwards to septum 4/5, however, in some species slightly beyond.
Segment – the primary unit of segmentation; a portion of the body internally separated
by septa, externally by intersegmental furrows. In most South African acanthodriline
species segmentation corresponds to external and internal transverse metameric
divisions, in clear distinction to indigenous microchaetids, where external annulation
is not congruent with internal divisions.
Seminal grooves – external elongated depressions connecting male pores with prostatic
pores.
Seminal vesicle (pl. seminal vesicles) – a part of male reproductive organs; a pocket
formed from dilated septum, where the sperm is matured.
Septum (pl. septa) – an internal transverse wall between segments, acting as a supporting
membrane for internal organs; usually delicate, although in some anterior segments
thickened to varying degrees. In the South African species its nature is a supportive
taxonomic character.
292 AFRICAN INVERTEBRATES, VOL 45, 2004
Seta (pl. setae) – any chitinous bristle, being a product of a single ectodermal cell at the
end of a tubular epidermal ingrowth. Setal formula is expressed in small letters:
a = the most ventral seta; b = the seta next lateral to a; c = the seta next lateral to b;
d = the seta next lateral to c; aa = median space ventrally between the two setae;
ab = space between a and b; bc = space between b and c setae; cd = space between
c and d setae; dd = space between the most dorsal setae. In the vast majority of
the endemic South African acanthodrilines setae are widely paired; it is not
confirmed whether the inter-setal ratio, often being variable, can be applied for
taxonomic evaluation.
Spermatheca (pl. spermathecae) – an organ in which sperm received from a copulatory
partner is stored. Each consists of the ampulla, duct, and diverticulum. The ampulla,
usually largest part of the spermatheca, is more or less ovoid, and thin-walled. The
duct is variable in shape and size, communicates by its ental part with the ampulla,
and through its ectal part with spermiductal pore. The diverticulum can be simple
or bi- or multiple-branched, variously shaped; its size and shape are of great
taxonomic importance. In South African endemic acanthodrilines, sperm is usually
stored in the diverticulum, sometimes also partly in the duct, but has not been
observed in the ampulla. The number, position and shape of spermathecae are
primary taxonomic characters.
Spermathecal pores – external openings at ends of ectal parts of spermathecae. Their
number and position are of high taxonomical importance.
Spermiductal funnel = testes funnel (pl. spermiductal funnels) – the ental part of sperm
duct through which sperm pass from testes into lumen of vas deferens on the way
to male pore.
U – circumference of the body; in South African Acanthodrilinae the distance between
dd is variable.
Vas deferens (pl. vasa deferentia) – the sperm duct carrying the sperm from the
spermiductal funnel towards the male pore.
Vesiculate – a holoic single nephridium, variable in size and shape of dilation forming
terminal bladder (a vesicle). A term used by Pickford (1937) for nephridia observed
in the species of Chilota and Microscolex.
TAXONOMY
Key to genera of the subfamily Acanthodrilinae known in South Africa
1 Holandric..............................................................................................................2
– Proandric...............................................................................................................3
2 Excretory system holoic, avesiculate. Male reproductive organs acanthodriline. Two
pairs of spermathecae; two pairs of spermathecal pores; each pair in intersegmental
furrow 7/8 and 8/9. One pair of male pores in segment 18, or in intersegmental
furrow 17/18. Two pairs of prostatic pores; each pair in segment 17 and 19. Gizzard
in segment 5; rudimentary, moderately or well developed. Spermathecal, prostatic
pores and male pores not approximate towards the mid-ventral line.....................
Eodriloides Zicsi, 1998
– Excretory system holoic, vesiculate. Male reproductive organs microscolecine,
variable. One or two pairs of spermathecae, or spermathecae absent; one pair of
spermathecal pores in intersegmental furrow 7/8 or 8/9, or two pairs, each in segment
PLISKO: UDEINA IN SOUTH AFRICA 293
7/8 and 8/9, sometimes absent. One pair of male pores in segment 17 or 18, or
absent. One pair of prostatic pores in segment 17, or two pairs, each in 17 and 19.
Male pores might be fused with prostatic pores or enter independently. Gizzard in
segment 5, rudimentary. In introduced species, a posterior pair of spermiductal
funnels might be reduced, or occurs in rudimentary parts. Spermathecal, prostatic
pores and male pores not approximate towards the mid-ventral line.....................
Microscolex Rosa, 1887
3 Excretory system holoic, vesiculate. Male reproductive organs acanthodriline. Two
pairs of spermathecae. Two pairs of spermathecal pores; each pair in intersegmental
furrow 7/8 and 8/9. One pair of male pores; male pores in segment 18. Two pairs of
prostatic pores; a pair in segment 17 and 19. Gizzard reduced, if present in segment
5, rudimentary or well developed. Spermathecal, prostatic pores and male pores
not approximate towards the mid-ventral line.............. Chilota Michaelsen, 1899
– Excretory system holoic, avesiculate....................................................................4
4 Male reproductive organs acanthodriline. Spermathecal, prostatic pores and male
pores not approximate towards the mid-ventral line, or situated close to the mid-
ventral line. Two pairs of spermathecae. Two pairs of spermathecal pores; each
pair in intersegmental furrow 7/8 and 8/9, or on the anterior border of segment 8
and 9. One pair of male pores in segment 18 or 19. Two pairs of prostates; two pairs
of prostatic pores, each pair in segment 17 and 19, or in 18 and 20. Gizzard in
segment 5 or 6; rudimentary, or moderately or well developed.............................
Parachilota Pickford, 1937
– Male reproductive organs balantine. Spermathecal, prostatic pores and male pores
not approximating towards mid-ventral line. One pair of spermathecae in 8 or 9.
Spermathecal pores in 7/8 or 8/9. One pair of male pores in segment 16 or 17 or 18,
or in intersegmental furrow 17/18; in front or behind of prostatic pores, separated,
or fused with prostatic pores. One pair of prostatic pores in segment 17 or 18 or 19.
One pair of well developed prostates, or one pair of well developed prostates and
rudimentary prostatic ducts in segment 17....................Udeina Michaelsen, 1910
Udeina Michaelsen, 1910 emend.
Holoscolex Ude, 1905: 421 (preoccupied by Cognetti 1904).
Yagansia [partim]: Michaelsen 1899a: 237.
Udeina Michaelsen, 1910: 53, 1912: 4, 5; Stephenson 1930: 671, 828; Pickford 1937: 416; Gates 1959:
239; Reynolds & Cook 1976: 62.
Udeina [applied for Parachilota]: Ljungström 1969: 370.
Udeina [partim]: Ljungström 1972: 100.
Udeina Michaelsen, 1890 [sic] Zicsi 1998: 67.
Type species: Yagansia kinbergi Michaelsen, 1899a: 443.
Revised diagnosis: Setae lumbricine. Prostomium tanylobous. Prostatic pores and male
pores not approximating towards the mid-ventral line. Prostates tubular. Proandric.
Holoic, avesiculate. One pair of spermathecae in segment 8 or 9. Spermathecal pores in
intersegmental furrow 7/8 or 8/9. One pair of male pores in segment 16 or 17 or 18 or in
17/18, in front or behind of prostatic pores, separated or fused with prostatic pores. One
pair of prostatic pores in segment 17 or 18 or 19. One pair of well-developed prostates,
or with additional vestigial parts of second prostatic pair. Spermathecal diverticulum
294 AFRICAN INVERTEBRATES, VOL 45, 2004
uni- or bilobate. Gizzard variably developed in segment 5. Typhlosole absent. Calciferous
glands absent.
Distribution: Endemic to South Africa. Known from Free State, North West province,
Western Cape, Eastern Cape, KZN.
Remarks and historical notes: The first species described with balantine features was
Yagansia kinbergi, collected over 150 years ago; the type locality was cited as ‘Caffraria’
or ‘Kaffraria’, possibly a part of western KZN extending into eastern Free State. A
single specimen, not clitellate, broken in two pieces, was described by Michaelsen
(1899a); subsequently it was restudied by Michaelsen (1907, 1910, 1912), and by Ude
(1905) who described the new genus Holoscolex for the new species reichei. Ude (1905)
compared these two species, both similarly characterised by balantine male reproductive
organs. However, no final clarification of the exact position of the male and prostatic
pores was stated. Michaelsen (1910), accepting the generic status suggested by Ude for
balantine species known from South Africa, proposed the new generic name Udeina
for Holoscolex because it was preoccupied. Subsequently, Michaelsen (1912)
synonymised reichei under kinbergi, stating: ‘Gen. Holoscolex Ude: da dieser Name
von Cognetti schon eine Glossoscolecinen-Gattung vergeben ist, so bedarf die Udesche
Acanthodrilinen-Gattung einer anderen Bezeichnung. Ich nenne sie Udeina, Typus:
U. kinbergi (Mich.) = Yagansia Kinbergi Mich. = Holoscolex Reichei Ude.’ To this,
Michaelsen also added information on the type locality, suggesting it was in the Free
State. In the same paper he wrote: ‘…Südafrika, etwa von der Breite der Kalahari
südwärts, charakterisiert durch die Microchätinen-Gattungen Microchaetus und
Tritogenia, sowie durch die Acanthodrilinen-Gattungen Chilota und Udeina’. Pickford
(1937) also synonymised U. reichei with U. kinbergi. Zicsi (1998), however, considered
that U. reichei may be a valid species, based on examination of material collected by
himself in the area of Parys on a bank of the Vaal River in the Free State.
The third species with balantine features is Udeina montanus Pickford, 1937 described
from a small sample from the Langeberg range in the Western Cape, differing from
U. kinbergi in many characters (location of prostatic and male pores, fused or separated
prostatic and male ducts, number of hearts). This species was accepted as more primitive
than U. kinbergi. An independent evolution of both species from different Parachilota
species was suggested. The species added by Zicsi (1998), U. pliskoae and
U. stuckenbergi, and six new species described in this paper—U. adriani, U. anneae,
U. mapelane, U. nkandla, U. petrosi, and U. qudeni—have raised the number of balantine
species to eleven. All being proandric, with holoic avesiculate nephridia, having only
one pair of spermathecae and one pair of prostatic pores, belong evidently to one species-
group, with the balantine condition characterising Udeina. It has been found that the
position of the spermathecal pores, male pores, and prostatic pores, are variable within
the genus (Table 2). However, the reduction of one prostatic pair, either the anterior or
posterior one, is evidently associated with the reduction of one pair of spermathecae
and relocation of male pores. The prostatic pores can occur anteriorly or posteriorly to
male pores, or be on the same segment as male pores. The ectal parts of the vasa deferentia
can enter into the male pores separately, or be fused with ectal parts of prostatic pores.
The presence of vestigial parts of the second pair of prostatic ducts observed in
U. petrosi may suggest stages in prostate reduction. It is noteworthy that a reduction of
PLISKO: UDEINA IN SOUTH AFRICA 295
TABLE 2
Selected characters used for the Udeina species separation.
Sperma- Male pores Prostatic Last pair of
Species name thecal pores pores pores Intestine hearts Diverticulum
adriani 7/8 18 19 16 12 unilobate
anneae 7/8 18 19 17 13 unilobate
kinbergi 7/8 17 18 16 13 unilobate
mapelane 7/8 18 17 16 12 unilobate
montanus 7/8 18 19 17 12 unilobate
nkandla 7/8 17/18 17 16 12 bilobate
petrosi 7/8 18 19 16 12 bilobate
pliskoae 7/8 18 17 16 12 unilobate
qudeni 7/8 16 17 15 12 unilobate
reichei 7/8 17 18 17 13 unilobate
stuckenbergi 8/9 17 17 17 12 unilobate
one pair of prostates, or relocation of prostatic pores or male pores, are variable in the
species currently grouped in Udeina; however, this variability was noted in populations
of U. anneae, U. mapelane, U. montanus, U. petrosi, U. pliskoae, and U. stuckenbergi.
A similar reduction of the anterior prostatic pair to vestigial prostatic ducts, was observed
in three dissected specimens of U. petrosi.
Key to species of the genus Udeina Michaelsen, 1910
1 Spermathecal pores in intersegmental furrow 7/8................................................2
– Spermathecal pores in intersegmental furrow 8/9.........stuckenbergi Zicsi, 1998
2 Prostatic pores in segment 17 or 18......................................................................3
– Prostatic pores in segment 19; male pores in 18 ..................................................7
3 Prostatic pores in segment 17...............................................................................4
– Prostatic pores in segment 18; male pores in 17; last pair of hearts in 13..........10
4 Male pores in 16 or in 17/18.................................................................................5
– Male pores in 18 ...................................................................................................6
5 Male pores in segment 16; intestine commences in 15; last pair of hearts in 12 ...
qudeni sp. n.
– Male pores in intersegmental furrow 17/18; intestine commences in 16; last pair of
hearts in 12; spermathecal diverticulum bilobate........................... nkandla sp. n.
6 Spermiductal funnels free; spermathecal duct extended; spermathecal diverticulum
large, attached with duct below ampulla; intestine commences in segment 16; last
pair of hearts in 12........................................................................mapelane sp. n.
– Spermiductal funnels enclosed; spermathecal duct short; spermathecal diverticulum
attached to duct at its base; intestine commences in segment 16; last pair of hearts
in 12.......................................................................................pliskoae Zicsi, 1998
7 Intestine commences in 16 ...................................................................................8
– Intestine commences in 17 ...................................................................................9
296 AFRICAN INVERTEBRATES, VOL 45, 2004
8 one pair of prostates in segments 17–21; last pair of hearts in 12; spermathecal
diverticulum unilobate......................................................................adriani sp. n.
– One pair of prostates in segment 19, and additional vestigial parts of prostatic ducts
in segment 17; spermathecal diverticulum bilobate ..........................petrosi sp. n.
9 Last pair of hearts in segment 12; spermathecal diverticulum unilobate ...............
montanus Pickford, 1937
– Last pair of hearts in 13; spermathecal diverticulum unilobate........ anneae sp. n.
10 Intestine commences in 16 ...................................... kinbergi (Michaelsen, 1899)
– Intestine commences in 17; last pair of hearts in segment 13 ................................
reichei (Ude, 1905)
Udeina adriani sp. n.
Figs 1–3
Etymology: Named in honour of Dr Adrian J. Armstrong of Ezemvelo KZN Wildlife,
who supported research on Acanthodrilinae in KZN, and collected this species.
Type material: Holotype: clitellate NMSA/Olig.03622 KwaZulu-Natal: Drakensberg,
section of Giant’s Castle Game Reserve, Injasuthi [new spelling: Inijasuthi]
(29º06'50.671"S:29º26'33.834"E), opposite camp, hillside at approx. 1540 m at south-facing
slope, wooded grassland at edge of dry drainage in Protea savannah, 2–10 cm below surface,
among roots of medium-tall Themeda grass, 14 June 2002, A. J. Armstrong.
Figs 1–3. Udeina adriani sp. n. 1. Holotype. Right spermatheca, extended, 400 X. A = ampulla;
Dv = diverticulum. 2. Left prostate, 180 X. Pg = prostatic gland; Pd = prostatic duct; Ps = penial
seta. 3. Ectal parts of penial setae, 500 X.