Corresponding author: Pavel Stoev (
Academic editor: Robert Mesibov
The centipede genus
The lithobiid subfamily
The identity of
The aim of present paper is to put on record all North African material of
The earliest record of the genus
In 1892 Pocock recorded
The first and hitherto only record of the genus in Libya comes from
Unless stated otherwise, the material treated herein has been collected from Tunisia in March 2008 and March 2009 by N. Akkari, P. Stoev and H. Enghoff, and also in the course of individual excursions of N. Akkari to different regions of the country in the period 2003-2008. The material is preserved in 70% or 96% ethanol and is shared between the Field Museum of Natural History, Chicago (
Eleven specimens from 5 species were used for genetic examination of the divergence among species of the genus. Ten specimens that sample 4 species were barcoded in the context of a global campaign on Myriapoda initiated as a part of the ‘Barcode of Life’ project (iBOL WorkGroup 1.9 ‘Terrestrial surveillance’) (Appendix C
Lysis of the tissues was carried out in 50 µl volume of lysis buffer and proteinase K incubated at 56°C overnight. DNA extraction followed a standard automated protocol on 96-well glass fibre plates (
The present paper demonstrates several innovative methods of semantic tagging and semantic enhancements, text and data processing, publishing and dissemination in taxonomy, described in more detail in a forum paper published in the same issue (
All 70 images included in this publication have been deposited in MorphBank (Appendix D
OD – original description; RD – redescription; K – key; FR – faunistic record; CHL – checklist or catalogue; BD – biological data.
T/TT – Tergite/Tergites, C – Coxa, Tr – Trochanter, PFe – Prefemur, Fe – Femur, Ti – Tibia; Letters a, m, p stand for spines in anterior, medial and posterior positions, respectively; those in brackets indicate the variable spines.
LI, II, III, IV stand for larval stadia 1, 2, 3, 4, respectively. PLI, II, III, etc. stand for post-larval stadia 1, 2, 3, etc. subad. = subadult; juv. – juvenile.
Order
Family
Subfamily
Medium- to large-sized
Several taxa assigned to
Type material of
Nontype material: SPAIN: 3 ♂♂, 3 ♀♀, 5 subad. ♂♂, 3 subad. ♀♀, 2 juv., labelled “
Length (from anterior margin of cephalic plate to posterior margin of telson) approx. 48 mm in largest specimens; cephalic plate slightly broader than long (
Cephalic plate finely punctate, with scattered minute setae, emerging from larger pits that form the punctae; slightly narrower than T1 (
Antennae moderately long, approx. 15 mm, reaching midline of T6 when folded backwards; approx. 37% length of body, composed of 43–44 articles; first three articles enlarged, with second being the largest (
Clypeus (
Forcipule (
Tergites wrinkled (
Sternites 1–14 elongated, subtrapeziform, finely punctate, with very few sparse setae, posterior margin convex (
Legs: all legs moderately long (
Coxal pores: small, circular, more concentrated on the outer part of pore-field, forming 3–4 irregular rows; only 2–3 pores from the internal row on each coxa larger; around 25–30 on legs 12–14 and about 17 pores on leg 15; pores of inner rows often separated by more than twice their own diameter, those of outermost row usually separated by less than their own diameter (
Male first genital sternite with emarginated posterior margin (
Female gonopods with 2+2 moderately long, apically pointed spurs and a simple, falcate claw (
Leg | Ventral | Dorsal | ||||||||
---|---|---|---|---|---|---|---|---|---|---|
C | Tr | PFe | Fe | Ti | C | Tr | PFe | Fe | Ti | |
1 | - | - | mp | amp | amp | - | - | amp | a-p | a |
2 | - | - | mp | amp | amp | - | - | amp | a-p | a-p\ |
3 | - | - | mp | amp | amp | - | - | amp | a-p | a-p\ |
4 | - | - | mp | amp | amp | - | - | amp | a-p | a-p\ |
5 | - | - | mp | amp | amp | - | - | amp | a-p | a-p\ |
6 | - | - | mp | amp | amp | - | - | amp | a-p | a-p\ |
7 | - | - | mp | amp | amp | - | - | amp | a-p | a-p\ |
8 | - | - | mp | amp | amp | - | - | amp | a-p | a-p\ |
9 | - | - | mp | amp | amp | - | - | amp | a-p | a-p\ |
10 | - | - | amp | amp | amp | - | - | amp | a-p | a-p\ |
11 | - | - | mp | amp | amp | - | - | amp | a-p | a-p\ |
12 | - | m | amp | amp | amp | a | - | amp | (a)-p | a-p |
13 | - | m | amp | amp | amp | a | - | p | (a)-p | |
14 | a | m | amp | amp | a-p | a | - | p | p | |
15 | a | m | amp | am | - | a | - | (a)m | p | - |
- “Pullus” with 10 pairs of legs + 2 incompletely developed pairs (= LIII)
- “Pullus” with 12 pairs of legs + 3 incompletely developed pairs (= LIV)
- “Juvenis” (smaller PL)
- “Junior” (larger PL)
His information is summarized in
Character states in larval and postlarval stadia in
Stadium | Body length (mm) | Number of antennal articles | Number of ocelli | Number of forcipular coxosternal teeth |
---|---|---|---|---|
Pullus (LIII) | 5-9 | 21 | 3 | 5+5 (also 4+5-4+4) |
Pullus (LIV) | 24-26 | 4 | ||
Juvenis | 9.5-11.8 | 25-29 | 4-6 | 4+5, 5+5 |
Junior | 13.5-20 | 35-42 | 6-8 | 5+5, 6+6 |
Character states in larval and postlarval stadia in
Stadium | Number of leg-pairs | Body length (mm) | Number of antennal articles |
---|---|---|---|
L0 | 7 | 5-6 | 9 |
LI | 8 | 6.5 | 11-13 |
LII | 9 | 7 | 15 |
LIII | 11 | 7.5 | 17 |
LIV | 13-14 | 8 | 21 |
PLI | 15 | 9-11 | 34-36 |
PLII | 15 | 13-15 | 38 |
PLIII | 15 | 16-18 | 38-39 |
PLIV | 15 | 19 | 39-40 |
PLV | 15 | 21-23 | 40 |
PLVI | 15 | 27-30 | 41-42 |
>PLVI | 15 | 33-45 | 42-43 |
Further data on the post-embryonic development based on Tunisian specimens of
In
Character states in larval and postlarval stadia in
Stadium | Number of leg pairs | Body length (mm) | Number of antennal articles | Number of ocelli | Number of forcipular coxosternal teeth | Sex | Locality |
---|---|---|---|---|---|---|---|
LIV | 12 | 7 | 26 | 3 | 4+4 | Gafsa, Jebel Bou Ramli | |
12 | 8 | 25 | 4 | 4+4 | Archeological site of Dugga | ||
12 | 9 | 25 | 5 | 4+4 | 6 km from Ouslatia | ||
PLI | 15 | 8 | 30 | 4 | 4+4 | Mides | |
PLII | 15 | 10 | 30 | 5 | 5+5 | 6 km from Ouslatia | |
PLIII | 15 | 14 | 41 | 7-8 | 5+5 | FF | Chambi National Park |
15 | 15 | 38 | 6-7 | 5+5 | MM | Jebel Bou Kornine | |
15 | 16 | 37-38 | 7-8 | 5+5 | MM | Archeological site of Dugga | |
15 | 17 | - | 7 | 5+5 | MM | Jebel Bou Kornine | |
15 | 18 | 44-46 | 7-8 | 6+6 | MM | Chambi National Park | |
PLIV | 15 | 18 | 44-46 | 10-411 | 6+6 | FF | Chambi National Park |
23 | 44 | 10 | 5+5 | MM | Tabarka* | ||
23 | 40 | 8 | 5+5 | FF | Tabrka* | ||
23 | 46 | 9 | 6+6 | FF | Tabarka* | ||
15 | 24 | 40-46 | 11 | 5+6 | FF | Beni Mtir | |
25 | 44 | 10 | 6+6 | FF | Tunis* | ||
15 | 26 | 45 | 11 | 5+5 | MM | Ghar El Melh | |
27 | 42 | 11 | 6+5 | MM | Tunis* | ||
28 | 41 | 13 | 6+6 | FF | Souk el Araba=Jendouba | ||
28 | 44 | 10 | 5+5 | MM | Tunis* | ||
PLV | 15 | 28 | 41-44 | 12 | 5+5 | MM | Dar Chichou |
15 | 25 | 41-42 | 14 | 6+6 | MM | Chambi National Park | |
29 | 44 | 12 | 5+5 | MM | Souk el Arba=Jendouba* | ||
30 | 43 | 13 | 5+5 | MM | Babouch* | ||
PLVI | 15 | 30 | 45-47 | 13-14 | 6+5 | MM | Ghar El Melh |
35 | 44 | 10 | 6+5 | MM | Souk el Arba=Jendouba* | ||
15 | 35 | 40 | 12-14 | 6+6 | MM | Beni Mtir | |
15 | 38 | 42-43 | 13-14 | 5+5 | FF | Beja | |
40 | 41 | 10 | 6+6 | FF | Tunis* | ||
15 | 40 | - | 14-15 | 5+5 | MM | Ghar El Melh | |
15 | 42 | - | 11-13 | 5+5 | MM | Jebel Bou Kornine | |
15 | 43 | 44 | 13 | 6+6 | MM | Sidi Khalifa | |
15 | 48 | 40-43 | 11-13 | 7+7 | MM | Sidi Khalifa |
Distribution of
Known from sea level up to approx. 1500 m. In Italy
W-Mediterranean, according to the chorotype classification of the W-Palearctic fauna proposed by Vigna Taglianti et al. (
Holotype: adult ♂, North Tunisia, Zaghouan Governorate, Jebel Zaghouan, Gouffre (chasm) Sidi Bou Gabrine, 36°22.423'N, 10°06.328'E, alt. 642 m, under clay lump, 17.III.2008, P. Stoev leg. (
A species of
Holotype: Length (from anterior margin of cephalic plate to posterior margin of telson) approx. 30 mm; cephalic plate slightly broader than long (
Cephalic plate smooth, wider than T1 (
Left antenna long, approx. 27 mm, reaching or slightly surpassing posterior margin of T12 when folded backwards; 90% length of body, composed of 68 articles; right antenna damaged, composed of at least 34 articles; basal two articles enlarged (
Clypeus with a cluster of about 30–33 long to medium-sized setae situated at apex and near the lateral margin (
Forcipule (
Tergites (
Sternites smooth, subtrapeziform, with few sparse setae, mainly at lateral margins. Posterior margins straight, slightly convex only on sternites 1 and 15 (
Legs: all legs generally elongated (
Spinulation: as in
Coxal pores: generally large, round to ovoid; 15–20 on legs 12–13 and about 20–24 on legs 14 and 15; pores separated by less than their own diameter, forming 3–4 irregular rows (
Male first genital sternite subquadrate (
Juvenile: pale yellow-whitish, with 15 leg-pairs, most detached; antennae long, approx. 90% of body length, composed of 36–37 articles; ultimate article almost 2.5 times length of penultimate; 5 ocelli; forcipular coxosternite with 5+5 teeth, median diastema shallow, V-shaped; TT 9, 11, 13 with posterior triangular projections; coxal pores: 2,1,1,1.
Leg | Ventral | Dorsal | ||||||||
---|---|---|---|---|---|---|---|---|---|---|
C | Tr | PFe | Fe | Ti | C | Tr | PFe | Fe | Ti | |
1 | - | - | mp | amp | amp | - | - | amp | A | a |
2 | - | - | mp | amp | amp | - | - | amp | a-p | a |
3 | - | - | mp | amp | amp | - | - | amp | a-p | a-p |
4 | - | - | mp | amp | amp | - | - | amp | a-p | a-p |
5 | - | - | mp | amp | amp | - | - | amp | a-p | a-p |
6 | - | - | (a)mp | amp | amp | - | - | amp | a-p | a-p |
7 | - | - | (a)mp | amp | amp | - | - | amp | a-p | a-p |
8 | - | - | amp | amp | amp | - | - | amp | a-p | a-p |
9 | - | - | amp | amp | amp | - | - | amp | a-p | a-p |
10 | - | (m) | amp | amp | amp | - | - | amp | a-p | a-p |
11 | - | m | amp | amp | amp | - | - | amp | a-p | a-p |
12 | - | m | amp | amp | amp | - | - | amp | a-p | a-p |
13 | - | m | amp | amp | amp | - | - | amp | a-p | a-p |
14 | - | m | amp | amp | a | a | - | am | a-p | a-p |
15 | a | m | amp | am | a | a | - | am | P | - |
derives from the Arabic word kahf (ﻒﮫﮐ) meaning ‘cave’, and kahfi denotes ‘living in cave’.
With very few exceptions, Tunisian specimens of
In order to confirm the delineation of the new species
Specimens sequenced for COI and their BOLD and GenBank accession numbers.
Species | Locality | GenBank accession number | BOLD accession number |
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Turkey, Afyon Prov., near village of Akoren |
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Tunisia, Cap Bon Peninsula, 20 km from El Haouaria |
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Tunisia, Cap Bon Peninsula, 7 km from Menzel Bou Zelfa |
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Tunisia, Nabeul, plage El Mansoura |
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Tunisia, Nabeul, plage El Mansoura |
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Tunisia, Jebel Zaghouan, surroundings of the marabout Sidi Bou Gabrine |
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Tunisia, Jebel Zaghouan, surroundings of the marabout Sidi Bou Gabrine |
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Tunisia, Jebel Zaghouan, Gouffre Sidi Bou Gabrine |
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Bulgaria, Shumen City |
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Bulgaria, Shumen City |
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Italy, Fogliano Mt, near Viterbo, Lazio |
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Genetic distances between species within
1 | 2 | 3 | 4 | ||
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1 |
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2 |
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21.4 | |||
3 |
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23.98 | 16.61 | ||
4 |
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19.19 | 22.28 | 23.46 | |
5 |
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21.47 | 18.15 | 20.7 | 21.56 |
Neighbor joining tree (K2P) of 5 species of
No taxonomically significant differences were found between the syntype specimens of
Information on the post-embryonic development of species of
a – A view of the entrance of cave Sidi Bou Gabrine, Jebel Zaghouan. b – A view of Jebel Zaghouan, Zaghouan Governorate, NE Tunisia.
Localities of
Distribution of
Locality of
In Tunisia
Unlike
1 (10) | Ocelli (Figs k-1-2) or posterior triangular projections on tergites absent (Fig. k-3) | 2 |
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2 (5) | Ocelli absent (Fig. k-1), posterior triangular projections at least on TT 9, 11, 13 (Fig. k-4) | 3 |
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3 (4) | Forcipular coxosternite with 9+9-12+12 teeth (Fig. k-5), forcipular trochanteroprefemur unmodified, 15VCa and 15 DCa spines present, pretarsus of leg 15 with a single claw (Fig. k-6) | |
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4 (3) | Forcipular coxosternite with 13-14 teeth (Fig. k-7), forcipular trochanteroprefemur strongly swollen, 15VCa and 15 DCa spines absent, pretarsus of leg 15 with a principal claw and posterior accessory claw (Fig. k-8) | |
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5 (2) | Ocelli present (Fig. k-9), all tergites without posterior triangular projections (Fig. k-10) | 6 |
6 (7) | Pretarsus of leg 15 with accessory claw | |
7 (6) | Pretarsus of leg 15 without accessory claw | 8 |
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8 (9) | T1 much broader than head (Fig. k-11), deeply concave posteriorly; forcipular trochanteroprefemur with a dorso-lateral knob (Fig. k-12) | |
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9 (8) | T1 as broad or slightly broader than head (Fig. k-13), transverse posteriorly; forcipular trochanteroprefemur without a knob | |
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10 (1) | Ocelli present (Fig. k-14); posterior triangular projections present at least on TT 11 and 13 (Fig. k-15) | 11 |
11 (20) | VCm spine present on leg 15 (Fig. k-16) | 12 |
12 (13) | Six ill-defined, feebly pigmented ocelli in adults | |
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13 (12) | 10-25 pigmented ocelli in adults (Fig. k-17) | 14 |
14 (19) | Male leg 15 with a large rounded knob proximate of the middle of the caudal side of the prefemur (Fig. k-18) | 15 |
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15 (18) | Antennae composed of 74-83 antennal articles | 16 |
16 (17) | Prefemoral knob simple (Fig. k-19) | |
17 (16) | Prefemoral knob apically incised forming two rounded processes densely covered with trichoid setae (Fig. k-20) | |
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18 (15) | Antennae composed of 50-60 antennal articles | |
19 (14) | Male leg 15 without prefemoral knob (Fig. k-21) | |
20 (11) | VCm spine absent (Fig. k-22) | 21 |
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21 (28) | Posterior triangular projections present on TT 9, 11, 13 or 11, 13 (Fig. k-23) | 22 |
22 (25) | Male gonopods long (Fig. k-24) | 23 |
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23 (24) | 17 ocelli, last 15 antennal articles shorter, only the ultimate article being 3 times longer than broad, others 2 times at most | |
24 (23) | Six-seven ocelli; last 15 articles of antennae elongated, 3 times longer than broad | |
25 (22) | Male gonopods short (Fig. k-25) | 26 |
26 (27) | Leg 15 approx. 30% length of body; prefemur of leg 15 in male moderately enlarged with two paramedian sulci not extending to posterior margin; posterior part of prefemur swollen dorso-medially (Fig. k-26); posterior margin of male first genital sternite emarginated (Fig. k-22) | |
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27 (26) | Leg 15 approx. 60% length of body; male prefemur 15 with a long, conical dorso-median protuberance (Fig. k-27); posterior margin of male first genital sternite not emarginated (Fig. k-28) | |
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28 (21) | Posterior triangular projections present on TT 6, 7, 9, 11, 13 or 7, 9, 11, 13 (Fig. k-29) | 29 |
29 (32) | Male gonopods short, two-jointed (Fig. k-25), VCa spine on leg 15 present (Fig. k-30), pretarsus of leg 15 without accessory claw | 30 |
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30 (31) | Prefemur of male leg 15 inflated and strongly expanded medially just proximad of the middle, the protuberance densely covered with setae (Fig. k-31); a densely setose circular dorsomedial area covering almost ? of prefemoral breadth in the position of DPFep spine, which is absent (Fig. k-31) | |
31 (30) | Prefemur of male leg 15 without such protuberance, evenly expanded along its whole length (Fig. k-32); circular area smaller, covering 1/3 of prefemoral breadth at most (Fig. k-32) | |
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32 (29) | Male gonopods long (unknown in |
33 |
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33 (34) | Antennae long, composed of 58-90 articles (Fig. k-34); leg 15 almost as long as body, densely covered with long setae 4-4.5 times diameter of article (Fig. k-35) | |
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34 (33) | Antennae and legs shorter (Fig. k-36), setae on leg 15 shorter (Fig. k-37) | 35 |
35 (42) | Pretarsus of leg 15 with accessory apical claw (Fig. k-33) | 36 |
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36 (39) | Seriate setae on tarsus 2 of leg 15 absent (Fig. k-38) | 37 |
37 (38) | Antennae and leg-pair 15 elongated, about 3/4 body length (Fig. k-36); femur of male leg 15 without basal pit (Fig. k-39) | |
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38 (37) | Antennae and leg-pair 15 about half body length; femur of male leg 15 with an extensive and deep basal pit (Fig. k-40) | |
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39 (36) | Seriate setae on tarsus 2 of leg 15 present (Fig. k-33) | 40 |
40 (41) | Basal pit of femur in male leg 15 extensive and deep; internal dorsal sulcus of femur in male leg 15 not extending to margin of pore-free area which bears a prominent globular swelling (Fig. k-41) | |
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41 (40) | Basal pit of femur in male leg 15 small and shallow; internal dorsal sulcus of femur in male leg 15 extending to margin of pore-free area which is not swollen (Fig. k-42) | |
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42 (35) | Pretarsus of leg 15 without accessory apical claw (Fig. k-43) | 43 |
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43 (44) | Pretarsus of leg 15 with a small accessory claw emerging basally to the principal claw (Fig. k-44), adults: 16-25 mm; T6 without posterior triangular projections (Fig. k-45) | |
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44 (43) | Pretarsus of leg 15 without accessory basal claw (Fig. k-43), body length of adults more than 30 mm; T6 with broad posterior triangular projections (Fig. k-46) | |
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The collecting trip of N. Akkari and P. Stoev in Tunisia in March 2008 was financially supported by the Field Museum Collection Fund, with the logistic help of Petra Sierwald. Atef Belkahla (Tunis, Tunisia) kindly helped with the preparation of maps, while Dicky Sick Ki Yu (Lexington, USA) with the exporting of the interactive key from Delta to Intkey format. Verena Stagl (NHMW) helped to obtain photographs of the type specimens of
Locality dataset for all published and new records of
Character dataset for larval and post-larval stadia of
BOLD dataset comprising the COI registration numbers of sequenced
MorphBank dataset of all published figures comprising figure number, species name, locality data, body part and additional metadata. File format: Microsoft Excel (1997-2003).
GoogleEarth (
Interactive key for identification of all currently valid species of
Copyright notice: All datasets are published under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided that the original author and source are credited.