Corresponding author: Uwe Jueg (
Academic editor: A. Schmidt-Rhaesa
Two new leech species from Africa are presented. The position of the eyes, the number of crop caeca and the gonopores separated by two annuli indicate that both belong to the genus
Eleven species of the genus
In February 2010, the malacologist Dr. Ulrich Bößneck with his wife Dr. Kathrin Bößneck (Nordhausen) and Dr. Christian Albrecht (University of Giessen) travelled to East Africa to search for small clams (
In Africa, several high mountain ranges with Afroalpine zones, some with glacial formation, stretch along the East African Rift Valley, from the Ethiopian highlands through Kenya and Uganda to Tanzania. Little is known about the leech fauna of these high mountains. The species newly described here should serve as a basis for further investigations on the leech diversity in these areas, which would likely result in the discovery of additional new species. The alpine enclaves of the high East African mountains offer a series of geographically and ecologically isolated temperate islands populated by a species-poor flora that is particularly adapted to the extreme climate (
All East African high mountains, e.g., Rwenzori Mountains (Moon Mountains), Kilamandjaro Massif, and Mt. Elgon Massif, show similar ecological conditions, which will be presented here using the example of Mt. Kenya Massif. The Mount Kenya massif, located about 150 km north of Nairobi, has the second highest mountain in Africa with the Batian (5,199 m above sea level). Its central areas, which are higher than 3.200 m, are part of Mount Kenya National Park. The massif rises within the Kenyan steppe as an enclave with special ecosystems. In the west, where the clouds accumulate, there is a tropical climate with a narrow belt of rainforest. The tree line is reached at approx. 3,200 m above sea level. There follows a zone up to approx. 4,000 m above sea level, which is mainly formed by tussock grasses, cushion plants and stemless rosette plants. Above this, the vegetation becomes increasingly sparse until finally, from 4,700 m above sea level, rugged rocks with no vegetation and scree slopes and glaciers follow. There are numerous mountain lakes, streams and waterfalls throughout the massif which drain into the Indian Ocean. The climate in the Afroalpine zone is characterized by extremes, with fluctuations of 20–30 °C observed within a day.
The genus
Flat glossiphoniid leeches with at least three pairs of eyes. The first pair is always close together and often fused. The following two pairs of eyes may be separated or compound. Salivary glands diffuse, six or seven pairs of crop caeca. Genital pores separated by 0–2 annuli. Cocoons or single eggs are attached to the venter (
Initially, there was only one specimen of the new species
A total of four species were hidden among these animals, including specimens of the new species
In addition to the comprehensive photographic representation of ten of the eleven known African species, complete distribution maps based on the current state of knowledge can also be presented for the first time for all species of the genus. In particular, the zoologists and photographers from the Zoological Museum of the University of Greifswald (Dr. Peter Michalik) and the Royal Museum of Central Africa in Tervuren, Belgium (Arnaud Henrard) created extensive photographic material of all known species (except
The following abbreviations are intended to simplify the naming of museums with more extensive material in the following text.
Anatomical features in the specimens examined could mostly not be described without damage to the animals. Nevertheless, clearly recognizable external features point to the current determinations, especially in the new species, which justifies their description. The holotype of
For the future, additional studies on the anatomy of the two newly described species are desirable, especially on fresh material. DNA sequencing is desirable for all species of the genus in order to clarify the relationships and the intraspecific range of variation of some species (e.g.
In addition to the holotype from Mt. Kenya, there were 20 specimens from the Mt. Elgon massif in Kenya, on the border with Uganda, which were collected in 1953 (
Metric characteristics of
Specimen-No. | Length | Maximum width | Length cranial sucker | Width cranial sucker | Length caudal sucker | Width caudal sucker | Length head region | Notes |
---|---|---|---|---|---|---|---|---|
1 holotype | 6.0 | 1.6 | 0.3 | 0.4 | 0.55 | 0.6 | 0.5 | |
2 | 7.5 | 2.6 | 0.45 | 0.5 | 0.75 | 0.75 | 0.38 | |
3 | 10.50 | 2.10 | 0.62 | 0.55 | 0.62 | 0.75 | 0.45 | |
4 | 10.50 | 2.00 | 0.52 | 0.60 | 0.45 | 0.60 | 0.40 | |
5 | 10.40 | 1.80 | 0.38 | 0.60 | 0.62 | 0.62 | 0.55 | |
6 | 9.70 | 1.35 | 0.55 | 0.55 | 0.55 | 0.60 | 0.50 | |
7 | 9.50 | 1.40 | 0.50 | 0.53 | 0.50 | 0.60 | 0.50 | |
8 | 9.20 | 1.20 | 0.35 | 0.48 | 0.48 | 0.50 | 0.50 | Eye position not visible |
9 | 9.10 | 0.90 | 0.50 | 0.50 | 0.42 | 0.42 | 0.48 | |
10 | 9.00 | 1.80 | 0.50 | 0.63 | 0.60 | 0.70 | 0.45 | |
11 | 8.30 | 2.20 | 0.50 | 0.65 | 0.75 | 0.80 | 0.50 | |
12 | 8.20 | 1.40 | 0.50 | 0.50 | 0.60 | 0.65 | 0.38 | Eye position not visible |
13 | 8.00 | 1.25 | 0.38 | 0.43 | 0.38 | 0.40 | 0.38 | |
14 | 6.00 | 1.90 | 0.53 | 0.55 | 0.55 | 0.60 | ? | head not removed |
15 | 5.70 | 1.30 | 0.60 | 0.65 | 0.75 | 0.75 | 0.40 | Eye position not visible; ventral glockenförmig (eggs) |
16 | 5.60 | 1.65 | 0.48 | 0.55 | 0.50 | 0.50 | 0.32 | |
17 | 5.30 | ? | 0.45 | 0.50 | 0.43 | 0.45 | 0.40 | atypically contracted |
18 | 5.30 | 1.30 | 0.53 | 0.53 | 0.50 | 0.50 | 0.38 | Eye position not visible; ventral glockenförmig (eggs) |
19 | 5.20 | 1.38 | 0.40 | 0.45 | 0.55 | 0.57 | 0.28 | Eye position not visible |
20 | 4.80 | 1.00 | 0.50 | 0.40 | 0.65 | 0.85 | 0.40 | ventrally bell-shaped (> 20 eggs) |
21 | 4.60 | 1.50 | 0.30 | 0.40 | 0.40 | 0.50 | 0.35 | |
Maximum | 10.50 | 2.20 | 0.62 | 0.65 | 0.75 | 0.85 | 0.55 | |
Minimum | 4.60 | 0.90 | 0.30 | 0.40 | 0.38 | 0.40 | 0.55 | |
Median | 7.54 | 1.58 | 0.48 | 0.53 | 0.56 | 0.61 | 0.43 |
1 specimen collected December of 1953 from Kenya, Mt.-Elgon-Massif near Kitale, 3,850 m above sea level; leg. Dr. J. Bouillon (
19 specimens collected December 11th, 1953 from Kenya, Mt.-Elgon-Massif near Kitale, east-northeast from the previous location, 3,000 m above sea level; leg. Dr. J. Bouillon (
Kenya, Central Kenya Province, Naro Moru, Mt. Kenya National Park north of the Teleki Valley, Nanyuki Tarn (mountain lake) (Fig.
Type locality of
The external morphology (number and position of eyes, annulation, colouration, papillation, suckers and the position of genital pores, size) was examined for the holotype and another 20 specimens. The internal morphology was not investigated (see above).
The newly described species is named after its occurrence in the Afroalpine zone of the East African mountain massifs Mt. Kenya and Mt. Elgon.
The affiliation to the genus
The gonopores of the species
Eye positions of
The geographic and habitat distribution distinguish
The mountain lake Nanyuki Tarn on Mt. Kenya (Fig.
Nothing is known about the ecology of the specimens found in 1953 in the Mt Elgon massif, except for the sea level at which the waters (probably mountain lakes) were located. Three of the animals carried eggs, so December can be named as the period for reproduction and brood care.
Distribution of
So far, the species is only known from the mountain massifs of Mt. Kenya and Mt. Elgon in Kenya. It is very likely that
A total of 25 specimens from two localities in Bunia in the Democratic Republic of Congo were examined, and are deposited in the Royal Museum of Central Africa in Tervuren (Belgium). They were labeled
Metric characteristics of
Specimen-No. | Length | Maximum width | Length cranial sucker | Width cranial sucker | Length caudal sucker | Width caudal sucker | Length head region | Notes |
---|---|---|---|---|---|---|---|---|
1 holotype | 7.10 | 3.00 | 0.25 | 0.40 | 0.95 | 0.95 | 1.60 | gonopores visible |
2 paratype | 6.70 | 2.55 | 0.25 | 0.15 | 0.75 | 0.75 | 1.38 | gonopores visible |
3 paratype | 6.50 | 3.15 | 0.25 | 0.25 | 0.75 | 0.83 | 1.40 | gonopores visible |
4 paratype | 6.10 | 2.30 | 0.20 | 0.15 | 0.70 | 0.65 | 0.75 | |
5 paratype | 6.50 | 2.15 | 0.25 | 0.35 | 0.65 | 0.70 | 1.38 | Gonopores faintly visible |
6 paratype | 5.90 | 2.25 | 0.30 | 0.30 | 0.70 | 0.68 | 1.00 | torn in the front part, Gonopores faintly visible |
7 paratype | 5.80 | 2.20 | 0.18 | 0.20 | 0.60 | 0.65 | 1.00 | |
8 paratype | 5.20 | 2.20 | 0.20 | 0.15 | 0.50 | 0.55 | 0.85 | |
9 paratype | 5.20 | 2.10 | 0.25 | 0.25 | 0.65 | 0.75 | 1.13 | |
10 paratype | 5.10 | 2.63 | 0.28 | 0.25 | 0.75 | 0.75 | 0.95 | Eyes clearly visible |
11 paratype | 5.10 | 2.00 | 0.23 | 0.15 | 0.60 | 0.58 | 0.95 | Crop caeca visible |
12 paratype | 5.00 | 2.75 | 0.20 | 0.20 | 0.63 | 0.65 | 1.20 | |
13 paratype | 5.00 | 2.50 | 0.20 | 0.15 | 0.62 | 0.65 | 1.30 | |
14 paratype | 4.50 | 2.05 | 0.25 | 0.20 | 0.50 | 0.50 | 0.85 | |
15 paratype | 4.13 | 1.75 | 0.20 | 0.15 | 0.40 | 0.48 | 0.68 | |
16 paratype | 4.75 | 1.90 | 0.18 | 0.15 | 0.50 | 0.50 | 0.75 | Crop caeca clearly visible against the light from anterior |
17 paratype | 4.70 | 1.95 | 0.20 | 0.18 | 0.50 | 0.50 | 0.85 | Crop caeca visible |
18 paratype | 4.60 | 2.65 | ? | ? | 0.50 | 0.55 | 0.65 | front end turned over |
19 paratype | 4.30 | 1.75 | 0.20 | 0.20 | 0.45 | 0.38 | 0.85 | Crop caeca visible |
20 paratype | 4.10 | 1.50 | 0.15 | 0.18 | 0.38 | 0.45 | 0.60 | |
21 | 4.60 | 2.80 | 0.25 | 0.30 | 0.62 | 0.50 | ? | Max. width in front of the middle, eyes not visible |
22 | 4.45 | 1.60 | 0.25 | 0.25 | 0.50 | 0.50 | ? | strong upper lip |
23 | 3–4 | 1.40 | 0.15 | 0.15 | 0.48 | 0.40 | ? | rolled up, eyes not visible |
24 | 3–4 | 1.15 | 0.10 | 0.10 | 0.38 | 0.35 | ? | rolled up, eyes not visible |
25 | 3–4 | 0.95 | 0.10 | 0.10 | 0.48 | 0.48 | ? | rolled up, eyes not visible |
Max | 7.10 | 3.15 | 0.30 | 0.40 | 0.95 | 0.95 | 1.60 | |
Min | 4.10 | 0.95 | 0.10 | 0.10 | 0.38 | 0.35 | 0.60 | |
Median | 5.32 | 2.17 | 0.21 | 0.21 | 0.59 | 0.60 | 1.03 |
Paratype 1: RMCA_VERMES_43663 (specimen 2 in Table
Paratype 2: RMCA_VERMES_43664 (specimen 3 in Table
Paratype 3: RMCA_VERMES_43665 (specimen 4 in Table
Paratype 4: RMCA_VERMES_43666 (specimen 5 in Table
Paratype 5: RMCA_VERMES_43667 (specimen 6 in Table
Paratype 6: RMCA_VERMES_43668 (specimen 7 in Table
Paratype 7: RMCA_VERMES_43669 (specimen 8 in Table
Paratype 8: RMCA_VERMES_43670 (specimen 9 in Table
Paratype 9: RMCA_VERMES_43671 (specimen 10 in Table
Paratype 10: RMCA_VERMES_43672 (specimen 11 in Table
Paratype 11: RMCA_VERMES_43673 (specimen 12 in Table
Paratype 12: RMCA_VERMES_43674 (specimen 13 in Table
Paratype 13: RMCA_VERMES_43675 (specimen 14 in Table
Paratype 14: RMCA_VERMES_43676 (specimen 15 in Table
Paratype 15: RMCA_VERMES_43677 (specimen 16 in Table
Paratype 16: RMCA_VERMES_43678 (specimen 17 in Table
Paratype 17: RMCA_VERMES_43679 (specimen 18 in Table
Paratype 18: RMCA_VERMES_43680 (specimen 19 in Table
Paratype 19: RMCA_VERMES_43681 (specimen 20 in Table
5 specimens; Democratic Republic of Congo, Ituri Province, Bunia, with
Democratic Republic of Congo, Ituri Province, Bunia, Jardin de Pères, near the border with Uganda at Lake Albert, leg. Courtois et Dr. Mass, 2nd March 1952 (in
A total of 25 specimens were examined, of which all external characteristics were recorded. Regarding the internal anatomy, the crop caeca could only be examined in backlight.
The newly described species is named after the place where it was found. Bunia is the capital of Ituri Province in the Democratic Republic of the Congo.
The placement in the genus
The gonopores of the species
Not much can be said about the ecology because no information other than the host animals was recorded when the specimens were collected in 1952.
Eye positions of
Distribution of
With the species newly described here, eleven representatives of the genus
The eastern Palaearctic
According to the distances between the gonopores, three groups can be distinguished worldwide within the genus
Two gonopores separated by 2 annuli:
Nine species belong to this group, of which only one species is found exclusively outside of Africa. This means that this trait is an original and almost unique trait of African species.
Two gonopores separated by 1 annulus:
Seven species can be classified in this group, their distribution areas are limited to Africa and Australia. Only the sole
One gonopore, common opening:
Nine species comprise this group. All species of the Holarctic (North America, Europe, Asia except Orientalis) belong here. The range of this group extends further into the area south of the Himalayas and even as far as Australia.
1 | Male and female gonopores united in one opening (basic color amber, only in NW-Africa) | |
– | Male and female gonopores separated by 1 or 2 annuli | 2 |
2 | Male and female gonopores separated by 1 annuli | 3 |
– | Male and female gonopores separated by 2 annuli | 4 |
3 | Crop caeca 6 (only in the Nile Delta) | |
– | Crop caeca 7 | |
4 | Body surface smooth, at most roughened by sensory papillae | 5 |
– | Body surface with distinct papillae or warts, usually arranged in distinct rows | 9 |
5 | Dorsally with distinct round to longitudinal oval spots on the annuli (Elongated body shape, only in the high mountains of Central Africa) | |
– | Dorsally unicolor or with faint longitudinal stripes (muscle strands) | 6 |
6 | Cranial sucker conspicuously pear-shaped (Edge of body strongly jagged, eyes fused (only 3 eyes visible), crop caeca 6, so far only in Central Africa) | |
– | Cranial sucker shaped differently | 7 |
7 | Mouth above center of cranial sucker (Basic color very variable (yellow, beige, blue, green, side edges hyaline, crop caeca 7, 200–400 eggs, in springs and spring streams, only Atlas Mountains) | |
– | Mouth in the center of cranial sucker | 8 |
8 | Dorsally irregular pattern of dark green chromatophores (Crop caeca 7, only in the lower reaches of the Nile) | |
– | Dorsally monochrome, faint longitudinal striations, lateral margins unpigmented (Crop caeca 7, widespread) | |
9 | Papillae irregularly arranged, body oblong (Brown ground color (dark even when fixed), crop caeca 7, only in the Nile) | |
– | Papillae arranged in (distinct) longitudinal rows | 10 |
10 | Proboscis almost as long as body, crop caeca 6, head not detached (Mouth with a conspicuous transverse slit, only found in a small area in South Africa) | |
– | Proboscis distinctly shorter than the body, crop caeca 7, head distinctly detached (Papillation very variable (possibly aggregated), almost throughout sub-Saharan Africa) |
Synonym:
The distribution area of
Distribution of
Earlier mentions of
Another Palearctic species that appears to be restricted to the south of the Iberian Peninsula (here the only species of the genus) and the Atlas Mountains is
Distribution of
The species
One specimen labeled
This species appears to be the rarest of the genus in Egypt. So far,
Distribution of
Synonyms:
The species was described from Lake Bunyoni in Uganda (
Only one voucher from the Central Africa Museum in Tervuren could be confirmed as
For Togo,
The species
Characterization of the type material used by
Specimen 1 | Specimen 2 | Specimen 3 | |
---|---|---|---|
|
oblong-oval; length 7,5 mm; maximum width 2,5 mm | oblong-oval; length 9,0 mm; maximum width 2,4 mm; Animal almost broken in the middle, front end turned over | oblong-oval; length ?; maximum width 3,0 mm; Animal folded up” several times |
|
indistinctly set off | indistinctly set off | indistinctly set off |
|
? (fixed light colorless) | ? (fixed light colorless) | ? (fixed light colorless) |
|
|||
|
dorsally many prominent papillae, per annulus at least 12, larger papillae on every third ring, arranged in 5–7 rows; ventrally smooth | dorsally many prominent papillae, per annulus at least 12, ventrally smooth | dorsally many prominent papillae, per annulus at least 12, ventrally smooth |
|
6, 1st pair close together, small, 2. and 3. pair fused left and right, much further apart, V-shaped | 6, 1st pair close together, small comma shaped, 2. and 3. Paar jeweils links und rechts schräg verschmolzen, deutlich weiter auseinander, breit V-förmig | 6, 1st pair confused, 2. and 3. pair closely together left and right (single eyes crescent-shaped), V-shaped |
|
small, extended proboscis (1,1 mm long); 0,25 × 0,35 mm | somewhat elongated/triangular, proboscis slightly extended; 0,20 × 0,20 mm | somewhat elongated/triangular, proboscis slightly extended (tip only); 0,35 × 0,38 mm |
|
in the middle, big | in the middle, slightly fissured | in the middle, slightly transverse |
|
circular, flat/even; 0,95 × 0,95 mm | circular, flat/even; 1,10 × 1,10 mm | circular, flat/even; dimensions ? |
|
7 (?), Postcaeca very well visible (with 4 simple lateral lobes) | 7 (?), 6 visible against the light (all unlobed), postcaeca not visible | ? |
|
2, separated by 2 annuli | ? (front end turned over) | 2, separated by 2 annuli |
Distribution of
Distribution of
According to the current state of knowledge, there is reliable evidence of
It would be worthwhile to study the intraspecific range of variation of different traits of
Distribution of
Synonym:
According to current knowledge,
Synonym:
Synonyms:
The species name of
Until now, the systematic classification of
The three animals were collected on May 13, 1911, by Prof. W. Michaelsen (Hamburg) in a pond on the Frauenstein farm near Neudamm (50 km ENE Windhoek, Namibia). Another species was found in the same pond,
Distribution of
According to
Distribution of
Characterization of the three types of
Lectotype | Paralectotype 1 | Paralectotype 2 | Total | |
---|---|---|---|---|
|
länglich-oval, oberseits gewölbt, ventral glockenförmig eingekrümmt; Länge 7,1 mm; max. Breite in 2. Körperhälfte 3,5 mm; Höhe 2 mm | ventral glockenförmig eingekrümmt (Individuum stark zusammengedrückt); Länge ca. 6 mm; max. Breite in 2. Körperhälfte ca. 3 mm, Höhe 2,5 mm | länglich-oval, oberseits gewölbt, ventral glockenförmig eingekrümmt; Länge 7,5 mm; max. Breite in 2. Körperhälfte 3,6 mm; Höhe 2,7 mm | länglich-oval, oberseits gewölbt, ventral glockenförmig eingekrümmt; Länge bis 7,5 mm; max. Breite in 2. Körperhälfte bis 3,6 mm; Höhe bis 2,7 mm |
|
short, set off by a nuchal furrow | short, set off by a nuchal furrow | short, clearly set off by a nuchal furrow | short, clearly set off by a nuchal furrow |
|
beige? | beige? | beige? | beige?; probably alive yellowish-brownish |
|
> 30 thin, very faint appearing dark, partly interrupted, longitudinal rows (clearly visible in the backlight); Edge transparent | > 30 thin, very faint appearing dark, partly interrupted, longitudinal rows (clearly visible in the backlight); Edge transparent | ca. 30 thin, very faint appearing dark, partly interrupted, longitudinal rows (clearly visible in the backlight); Edge transparent | 30 thin, very faint appearing dark, partly interrupted, longitudinal rows (clearly visible in the backlight); Edge transparent |
|
small flat papillae evenly distributed over the body; looks rather smooth, median slight increase recognizable (no crest) | none; Skin surface uneven due to sensory papillae, but no prominent papillae | small flat papillae evenly distributed over the body; looks rather smooth, sensory papillae equally present; median slight increase recognizable (no crest) | Prominent papillae absent, small flat papillae evenly distributed over the body; looks rather smooth, median slight increase recognizable (no crest) |
|
6; 1st pair close but separated, slightly shifted to the righ, on a4, distance between 1. and 2. pair greater than between 2. and 3. pair; 2. and 3. pair merged left and right respectively, on a6 | 6; 1st pair fused but separated, on a4; 2. and 3. pair each left and right fused, auf a6 | 6; 1st pair fused but separated, on a4; 2. and 3. pair each left and right fused, on a6 | 6; 1st pair fused, partially merged, on a4; 2. and 3. pair each left and right fused, on a6 |
|
roundish-acrossoval, moderately deep, thin walls; 0,8 × 0,9 mm | roundish-acrossoval, moderately deep, thin walls; 0,6 × 0,8 mm | oblong-oval, moderately deep, thin walls; 0,6 × 0,5 mm | roundish-acrossoval, moderately deep, thin walls; 0,6 × 0,7 mm |
|
in the centre, big | in the centre, slightly fissured | in the centre, slightly transverse | in the centre, different sized |
|
acrossoval, almost square, moderately thick walls; 1,3 ×1,0 mm; directed ventrally | oblongoval; 1,1 × ? mm (strongly compressed); directed ventrally | roundish; 1,0 × 1,0 mm; directed ventrally | oblongoval, moderately thick walls; 1,1 × 1,0 mm; directed ventrally; posterior edge barely visible from dorsal |
|
? | ? | ? | ? |
|
? (possibly 2, separated by 2 annuli) | ? | 2, separated by 2 annuli) | 2, separated by 2 annuli, very small, hard to see |
none | 30 Juveniles ventral | 26 eggs ventral | clutches with at least 30 eggs/juveniles |
*
A comparison of
Characteristics of African
|
oval to pear-shaped, broadly oval at rest, up to 10 mm long, up to 5 mm wide | elongated oval at rest, up to 16 mm long, up to 6 mm wide | oval to oblong when at rest, up to 13 mm long, up to 5 mm (?) wide | elongate-oval at rest, widest behind the middle, 7 mm long, 2.5 mm wide, dorsally convex, ventrally flat | elongate-oval, up to 8mm long, up to 3mm wide |
|
bluntly rounded | bluntly rounded | slightly pointed | not deposed, no narrowing of the neck | discontinued |
|
orange-amber, rarely yellowish to white-grey | blue, green, turquoise, beige, brownish, yellowish, head and sides transparent | light brown with dark brown chromatophores | snow-white, slightly transparent | greenish |
|
no | no | irregular dark brown longitudinal stripes | dark brown pigments on the tubercles of the central crest (on a2), irregularly also on other warts on a2 | irregularly arranged pattern of dark green chromatophores |
|
finely granulated by faint even papillae | smooth or small regularly arranged sensory papillae | 12–19 large papillae per annulus, irregularly arranged, many small papillae | Approx. 20 warts of unequal size per annulus, median row of warts like a crest, small papillae | smooth, without papillae except minute papillae in cranial sucker |
|
6, 1st pair mostly separate, 2nd and 3rd often fused, also absent | 6, 1st pair mostly fused, 2nd and 3rd often fused, also missing | 6, 1st pair fused, 2nd and 3rd pair with large space between and fused | 6, 1st pair mostly fused, 2nd and 3rd often fused, also missing | 6, 1st pair mostly fused, 2nd and 3rd often fused, various eye positions |
|
large, broad, bowl-shaped | pointed transversely oval, evenly curved in front, mostly with a blunt edge behind, filled in | Narrow transverse oval, evenly arched in front and behind, bowl-shaped | bowl-shaped with a thickened edge | cup-shaped, with a thickened rim |
|
small, in the center of cranial sucker | small, slightly above center of cranial sucker | small, in the center of cranial sucker | subapical position | small, in the center of cranial sucker |
|
large, ± circular, > than half the maximum body width | small, ± circular, < half the maximum body width | small, ± circular, < half the maximum body width | small, ± orbicular (?), < half maximum body width, narrow crescent-shaped margin visible dorsally | small, circular, < 1/3 of maximum body width |
|
6, ± evenly curved, only the posterior branched | 7, angled, all branched | 7, all branched | 6 | 7, bilobed |
|
1 | 2, separated by 2 annuli | 2, separated by 2 annuli | 1 | 2, separated by 2 annuli |
|
30–60 | 200–400 | 30–50 | ? | 13–25 |
|
|
|
|||
|
rivers, lakes, ponds | sources and streams in the colline and montane regions | streams, rivers and canals, on stones, never on plants | stream and channels, under stones and at the base of water lilies ( |
shallow, muddy, polluted calm water |
|
Western, Central and Eastern Europe, Balkans, Italy, not on the Iberian Peninsula, to Central Asia, Northwest Africa | Southern Iberian Peninsula, Morocco (Atlas Mountains) | Egypt (Nile between Luxor and Delta), Ethiopia (?) | Egypt (Nile and irrigation canals in the Nile Delta) | Egypt (Al Sont Canal in Asyut) |
Characteristics of African
|
elongated, moderately curved dorsally, flat ventrally, 6.0 mm long, 1.6 mm wide, posterior edge strongly concave dorsally (part of the caudal sucker is exposed) | elongated-oval, dorsally moderately convex, ventrally flat, up to 7.1 mm long, up to 3.1 mm wide, tapering towards the front end, with head lobes, lateral edges wavy to very jagged | elongate-oval, first third narrow, 11.6 mm long, 2.2 mm wide, sexual maturity already at a small size, slightly convex, flat underside | pointed-oval, tapering towards the front end, flattened, up to 9.8 mm long, up to 4.2 mm wide | elongate-oval, moderately flattened, ventrally flat, 9.1 mm long, 2.8 mm wide, sexually mature even at a small size | elongate oval, moderately flattened, dorsally convex, ventrally flat to bell-shaped, up to 12.5 × 2.8 mm |
|
distinct, short and wide | not clearly set off, tapered in front | clearly set off over neck constriction | not deposed or widened | clearly separated, only slightly broadened | short, clearly set off by a nuchal furrow |
|
probably light brown to beige | probably light brown to yellowish | flesh-colored or green | brown | green or brown, or shades thereof | brown or green |
|
circular spots (only slightly darker than ground colour) on each annulus, giving a diffuse pattern on the dorsal side, suggesting (probably coincidentally) longitudinal rows which may be interrupted or of different lengths, higher concentration of circular spots on the sides of the body, fewer ventrally stain | 30–36 narrow dark vertical stripes, no spots or patterns | dorsally with about 36 longitudinal stripes at the widest part of the body or chromatophores irregularly distributed | more than 10 pairs of dark brown narrow longitudinal stripes, light median field, isolated pigments outside the stripes | 34–36 dark longitudinal stripes at the widest part of the body; Chromatophores evenly distributed dorsally except for head region, lateral margins and posterior sucker, sometimes forming transverse streaks, paler unicolor ventrally | dorsally with 30 dark brown or dark green narrow discontinuous stripes, fainter towards the ends, apex, postnatal ring and lateral margins unpigmented |
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smooth, no warts or papillae, very small, barely visible, low sensory papillae in transverse rows on each annulus | smooth, no warts or papillae, very small, barely visible, low sensory papillae, especially numerous on the margins of the lateral lobes | 18 distinct papillae in transverse rows on each annulus in midbody region; larger papillae arranged in 7 longitudinal rows, many very sensory papillae unevenly distributed | roughened by clear papillae (largest on a2) on the dark longitudinal stripes | 3 pairs of longitudinal rows of papillae on both sides, 16 large papillae on each annulus, numerous small rounded papillae, ventrally fewer small papillae | no prominent papillae, very small, low dome-shaped sensory papillae evenly distributed over body, eye area and posterior suction cup with irregular papillae |
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6, 1st pair on a4, 2nd and 3rd pair on a5, eyes of 2nd and 3rd pair larger, occasionally fused, great variability in arrangement | 6, 1st pair (fused) on a4, 2nd and 3rd pair (left and right fused) on a5, only 3 eyes visible | 6, 1st pair on a4, 2nd and 3rd pair most fused or close, both on a5 | 6, 1st pair of smaller ones, anterior and both lateral pairs often fused, variable | 6, 1st pair on a4, 2nd and 3rd pair on a5, eyes of 2nd and 3rd pair much larger, occasionally fused | 6, small, 1st pair in a4, 2nd and 3rd in a5, pairs of eyes close, sometimes fused, variable |
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large, cupped, with strong edges | small, flat, pear-shaped, rounded in posterior part, grooved in front, thin-walled | small, shovel-shaped | flat | cupped with deep cavity, well-developed rim with numerous tiny papillae | roundish-oval, moderately deep, bowl-shaped with thin side walls, edge covered with small papillae; 0.6 × 0.7mm |
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in the center, mostly large, triangular, sides slightly concave, base slightly convex | in the center, stitch-like small | relatively large in the center | in the center, with a striking transverse slit | big in the center | large, on top of a large dome-shaped papilla, in the center |
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small, circular, < half of maximum body width, slightly cupped, deepened in the middle, with a broad margin (ring-shaped) | small, circular, < third of maximum body width, flat, broad base, with thick wall | small, circular, diameter 0.9 mm, < half of maximum body width | small, circular, < half maximum body width, flat cup-shaped, distinct from body, dorsally with 7–8 brown bands | small, circular, < half maximum body width, deeply cupped with thin margin, broadly attached | small, circular-elongated oval, 1.1 × 1.0 mm; moderately thick walls; Posterior edge barely visible from dorsal |
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? | 6, first 5 pairs slightly divided, 6th pair each with 5 small side lobes | 7, the first 6 unlobed, 7th pair each with 4 small side lobes | 6, first 5 pairs unlobed, 6th pair with 4 side lobes each | 7, bilobed, postcaeca with 5–6 lateral lobes | 7, the first 6 bilobed, postcaeca each with 15 slender lateral secondary lobes |
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2, separated by 2 annuli, very small and inconspicuous, areas visible only as narrow dark streaks in the sulci between the annuli | 2, separated by 2 annuli, very small and inconspicuous | 2, separated by 2 annuli (XI/XII and XIIa2/a3), male well visible, female not | 2, separated by 2 annuli, both small | 2, separated by 1 annulus, walls at male and female atria | 2, separated by 2 annuli, very small, poorly recognizable |
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clutch with at least 20 eggs | ? | ? | ? | ? | clutch with at least 30 eggs |
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probably small mussels ( |
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mountain lakes in the Afroalpine zone (3,000 to 4,500 m above sea level) | ? | permanent and semi-permanent bodies of water: rivers, lakes, reservoirs with muddy water, under stones | reservoirs, on riparian vegetation | springs with permanent water, rainwater pools, reservoirs with muddy water, under tree stumps | rivers (also ephemeral), rainwater basins, ponds, reservoirs, under stones |
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Kenya (Mt. Kenya Massif and Mt. Elgon Massif) | Democratic Republic of Congo, Bunia (Province of Ituri) | Uganda, Ethiopia, Togo, Democratic Republic of Congo, Namibia, South Africa | Coastal region of South Africa between Mozambique and Lesotho | Namibia, South Africa | Namibia, South Africa, Democratic Republic of Congo, Kenya |
The type material for
In the Africa Museum Tervuren (Belgium) under the coll. no. 558 et 564 there are 10 voucher specimens labeled
Distribution of
A specimen of this species was also found among the material in the Natural History Museum London, which was inventoried under
Thus,
I would like to thank Messrs. Christophe Allard and Dr. Didier Van den Spiegel (Royal Museum of Central Africa in Tervuren, Belgium) for sending extensive historical museum material from the Sciacchitano collection (
For the time-consuming photography of important species and specimens, a big thank you goes to Dr. Peter Michalik (Zoological Institute and Museum of the University of Greifswald) and to Arnaud Henrard (Royal Museum of Central Africa in Tervuren, Belgium). His images were taken in the framework of the Belgian Science Policy (BELSPO) project DIGIT04. I thank Brigitta and Frank Eiseler (Roetgen) for using a photo of
Dr. Michael L. Zettler (Leibniz Institute for Baltic Sea research Warnemünde), Christophe Allard (Royal Museum of Central Africa in Tervuren, Belgium) and Nasser El-Shiemy (University of Assiut, Egypt) helped with the literature procurement, for which they are gratefully acknowledged. Dr. Kathrin Bößneck (Nordhausen) I would like to thank for providing information on the locality of
Thanks to Mariam Dunker, Dr. Jenna Moore and Prof. Dr. Schmidt-Rhaesa (Zoological Museum Hamburg) for helping me with the professional translation of the text into English.