European Journal of Taxonomy 139: 1-11

http://dx.doi.org/10.5852/ejt.2015.139

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ISSN 2118-9773

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2015 • VandenSpiegel D. et al.

Research article

um:lsid:zoobank.org:pub:F7065B0E-FCCC-45EC-9C4F-19F6C9B972B3

Two new species, including one representing a new genus, of the West

African millipede family Campodesmidae (Diplopoda: Polydesmida)

D. VANDENSPIEGEL 1 , S.I. GOLOVATCH 2 & A.R. NZOKO FIEMAPONG 3

1 Musee Royal de l’Afrique Centrale, B-3480 Tervuren, Belgique.

Email: [email protected] (corresponding author)

2 Institute for Problems of Ecology and Evolution, Russian Academy of Sciences,

Leninsky prospekt 33, Moscow 119071, Russia.

Email: [email protected] (corresponding author)

3 Laboratoire de Zoologie, Universite Yaounde 1, BP812, Cameroon.

Email: [email protected]

1 um:lsid:zoobank.org:author:CE8C3D01-28AD-43F7-9D4F-04802E68CBlA

2 um:lsid:zoobank.org:author:71532F45-BDD5-415D-BC54-86256E5D5D4A

3 urn:lsid:zoobank.org:author:ECDC7821-D8DA-49D4-9D6A-13A6E732863E

Abstract. A new genus, Campodesmoides gen. nov., is described to only encompass C. corniger sp.

nov., from Cameroon. This genus and species is distinguished from the few known species of the small

western African family Campodesmidae, all currently in Campodesmus , by the much longer antennae

and legs, the normal pore formula with ozopores borne on porosteles, and the suberect and distally twisted

gonopod, coupled with peculiar horns on a few anterior postcollum segments. A new Campodesmus is

also described, C. alobatus sp. nov., from Ivory Coast, which differs from congeners primarily in the

lack of a dorsal/lateral lobe on the otherwise usual and strongly subcircular gonopod telopodite, albeit

the latter is not directed mesad, but held subparallel to the main body axis.

Key words. Myriapoda, taxonomy. Ivory Coast, Cameroon.

VandenSpiegel D., Golovatch S.I. & Nzoko Fiemapong A.R. 2015. Two new species, including one representing

a new genus, of the West African millipede family Campodesmidae (Diplopoda: Polydesmida). European Journal

of Taxonomy 139: 1-11. http://dx.doi.org/10.5852/eit.2Q15.139

Introduction

The small millipede family Campodesmidae is currently known to comprise “about eight species” from

Liberia, Sierra Leone, Ghana, Ivory Coast and Nigeria (Hoffman 1980, 1982), albeit Demange (1971)

had considered and keyed only seven. All of them are assigned to the genus Campodesmus Cook, 1896,

which is characterized by a dark medium-sized body (17-38 mm long) with more or less strongly

tuberculate metaterga and only moderately declivous paranota (Schiotz 1966). The modestly or even

not enlarged paranota 2 suggest incomplete or imperfect volvation, i.e., enrollment into only a partial

coil (Golovatch 2003). Ozopores are only present on paranota 5 and 7. The gonopod aperture is rather

large, transversely oval, the edges at least partly being elevated and the gonopods almost fully exposed.

European Journal of Taxonomy 139: 1-11 (2015)

The gonocoxae are elongated subcylindrical, fused basally, at best poorly setose, concealed inside the

aperture. The telopodites are long, strongly subcircular, directed anteromesally, crossing in situ , on the

dorsal/lateral face usually with a prominent, variously shaped lobe (= prefemoral process) at the base of

the telopodite.

Interestingly, the course of the seminal groove has never been traced and properly described in the

literature (Hoffman 1951, 1982; Schiotz 1966; Demange 1971). Based on the available illustrations,

however, the family and genus could additionally be characterized by the lack of a special solenomere

to support at least the terminal part of the seminal groove.

The present paper records two new species of Campodesmidae: 1) a new Campodesmus from Ivory

Coast, and 2) a second genus of the family, Campodesmoides gen. nov., a monotypic taxon from

Cameroon. These two genera are markedly distinct from each other both in gonopod conformation and

several somatic characters.

Material and methods

All material treated here derives from the collection of the Musee Royal de T Afrique Centrale (MRAC),

Tervuren, Belgium, with only one paratype shared with the holdings of the Zoological Museum of the

Moscow State University (ZMUM), Moscow, Russia. The samples are stored in 70% ethanol. Specimens

for scanning electron microscopy (SEM) were air-dried, mounted on aluminium stubs, coated with gold

and studied using a JEOL JSM-6480LV scanning electron microscope. Photographs were taken with a

Leica DFC 500 digital camera mounted on a Leica MZ16A stereo microscope. Images were processed

with the Leica Application Suite software.

Results

Class Diplopoda Blainville-Gervais, 1844

Order Polydesmida Leach, 1814

Family Campodesmidae Cook, 1895

Genus Campodesmoides gen. nov.

urn:lsid:zoobank.org:act:D7DFD21A-C64C-40E4-9CA2-637CAB4E16A2

Diagnosis

Same as for Campodesmoides corniger sp. nov., due to monotypy of this genus.

Etymology

The name of this genus emphasizes the attribution of the new genus to the Campodesmidae which has

been known so far by the single genus Campodesmus. Gender masculine.

Type species

Campodesmoides corniger sp. nov.

um:lsid:zoobank.org:act:E20A8CC8-F56C-4499-BlF0-F3D541275368

Figs 1-3

Diagnosis

A species of Campodesmidae which differs from other species of the genus Campodesmus , the only

hitherto recognized genus of Campodesmidae, by the long and only slightly clavate antennae, which

VANDENSPIEGEL D. et al.. Two new Campodesmidae

reach behind segment 3 when stretched dorsally; the considerably longer legs which in the male are

about 1.5 times as long as midbody height; the normal pore formula: 5, 7, 9, 10, 12, 13, 15-19, the

ozopores being borne on conspicuous porosteles (versus reduced to 5, 7 without porosteles); the more

dorsally situated paranota which are set at the upper % of midbody height (versus about half of midbody

height); the nearly flat sterna and the gonopod telopodites being suberect, in situ directed forward, not

crossing mesally, each with a clearly twisted solenophore devoid of a distinct solenomere.

Etymology

To emphasize the high, vertical, paramedian horns present on metatergum 2, these growing increasingly

small on metatergum 3 and, especially, 4; noun in apposition.

Material examined

Holotype

CAMEROON: Ongot, N 03°52’, E 011°2E, c. 830 m a.s.l., open terrain, 26 Apr. 2014, leg. A.

Nzoko, MRAC 22679.

Fig. 1 . Habitus of Campodesmoides corniger sp. nov., holotype, lateral view. Photo by J. Brecko, not

to scale.

European Journal of Taxonomy 139: 1-11 (2015)

Paratypes

CAMEROON: 1 §, same locality, together with holotype, MRAC 22.681; 1 same locality, 29 Nov.

2014, MRAC 22.678; 1 ?,Egoutadjap,N02°43 , ,E 11°04’, forest, 06 Dec. 2014, MRAC 22.680 (SEM).

All leg. A. Nzoko.

Description

Length of holotype c. 26 mm, width on midbody pro- and metazonae 2.5 and 5.0 mm, respectively,

female paratypes 25-30 mm long, 2.5-3.6 and 3.6-5.0 mm wide on pro- and metazonae, respectively.

Coloration from light grey-brown (female) to blackish brown (male, female) (Fig. 1), darker specimens

Fig. 2. SEM micrographs of Campodesmoides corniger sp. nov., $, paratype, MRAC 22.680. A, D,

G. Anterior part of body, lateral, dorsal and ventral views, respectively. B, E, H. Midbody segments,

lateral, dorsal and ventral views, respectively. C, F, I. Posterior part of body, lateral, dorsal and ventral

views, respectively. J. Cross-section of a midbody segment, caudal view. K. Limbus and adjacent

structures. L-M. Poriferous paratergite with porostele, dorsal and lateral views, respectively. Scale bars:

A-J = 0.5 mm : K-M = 0.1 mm

VANDENSPIEGEL D. et al.. Two new Campodesmidae

with brown to chocolate brown antennae and legs; tips of antennae, distal parts of paranota 5, 7, 9 and 13

together with their porosteles, as well as the remaining porosteles (male, female) or only all porosteles

(female) yellowish to nearly pallid.

Head. Except for a bare and rather densely setose clypeolabral region, very densely microdenticulate-

microtuberculate-microgranulate, interantennal isthmus almost as broad as diameter of antennal socket

(Fig. 2). Antennae long and only slightly clavate due to a somewhat higher antennomere 6, in situ

reaching behind segment 3 when stretched dorsally; only antennomere 6 with a dorso-apical group of

tiny bacilliform sensilla; in length, antennomere 6>2>3 = 5>1>7; apical segment with the usual

four sensory cones.

Body. With 20 segments (male, female). In width, segment 3 = 4 < head = collum = 5-17 < 2 (male),

or collum < head = 3 = 4<2<5-17 (female); body rapidly tapering from segment 18 towards telson.

Collum transversely ellipsoid, far from covering entire head from above, dorsal surface rather regularly

and densely tuberculate, with two transverse rows of higher 2 + 2 and 4 + 4 tubercles in caudal half,

paranota bearing 2-3 lateral teeth (Figs 1, 2A, D). Prozonae smooth and weakly shining; metazonae

dull, densely microtuberculate or microgranulate all over, cerotegument well-developed, sometimes

in places clothed with a crust of soil; dorsal surface usually with three regular, transverse rows of

normally small tubercles or short spines; stricture very delicately striolate. Metatergum 2 with two

transverse rows of spines, 2 + 2 per row, mid-dorsal pair in rear row being only slightly separated,

unusually high, vertical and horn-shaped compared to very small remaining spines; paranota 2 strongly

elevated, directed dorsolaterad at about 45°, each with 3-4 apical teeth (Figs 1, 2A, D). Tuberculation

pattern persisting on metaterga 3 and 4 as well, but mid-dorsal pair of horns first considerably and then

much shorter, respectively, each horn thereby carrying a subapical denticle on caudal face (Fig. 2A,

D); paranota 3-8(9) increasingly less strongly elevated to become subhorizontal (Figs 1, 2B-C, E-F,

J). Metaterga 5-19 with a faint, but visible, transverse, midtergal impression separating first transverse

row of 1 + 1 (segment 5), 2 + 2 (segments 6-14) or 3 + 3 (segments 15-19) small tubercles from two

further transverse rows, middle row usually consisting of slightly higher tubercles: 3+3 (segments

5-14) or 4(5) + 4(5) (segments 15-19); caudal row located near caudal margin, usually consisting of

Fig. 3. Campodesmoides corniger sp. nov., holotype. A-B. Feft gonopod, mesal and lateral views,

respectively. C-E. Right gonopod, mesal, lateral and ventral views, respectively. Scale bars = 0.2 mm.

Abbreviations are explained in the text.

European Journal of Taxonomy 139: 1-11 (2015)

particularly small tubercles, yet clear enough against a microgranulate/microtuberculate background:

2 + 2 (segments 5-14) or 3(4) + 3(4) (segments 15-19). Poreless paranota usually with 4-5 distinct

teeth at lateral margin, pore-bearing ones normally subtruncate, with 1-2 teeth at both fore and rear

corners, and a prominent, midlength porostele (Figs 1, 2B-C, E-F, J). Pore formula normal, ozopores

always borne on somewhat mushroom-shaped porosteles. Paranota set high (mostly at about upper %

of body height, dorsum thus being only moderately convex), inclined caudad and increasingly clearly

protruding behind rear tergal margin from segment 17 towards telson (Figs 1, 2C, F), considerably

broader, and porosteles longer in male than in female. A t hin dark axial line somet im es traceable through

a transparent tegument. Pleurosternal carinae absent, a few small tuberculations in their place (Fig.

2A). Limbus entire (Fig. 2K). Epiproct short, small, subtruncate, coniform, with a pair of prominent

subapical papillae (Fig. 2C, F, I). Hypoproct nearly semi-circular with 1 + 1 caudal setae very distinctly

separated and borne on minute knobs (Fig. 21).

Sterna. Broad, about twice as broad as coxa length, almost flat, axial impression absent (Fig. 2H).

Gonapophyses on male coxae 2 vestigial (Fig. 2G). Legs long to very long, about 0.9-1.0 or 1.5-

1.6 times as long as midbody height (female and male, respectively), very slender; in length, femur >

tarsus > prefemur = postfemur = tibia = coxa; claw very small, very slightly curved; ventral surface of

tarsi densely setose, but forming no brushes.

Gonopod. Aperture transversely ovoid, large, its lateral and posterior edges slightly elevated, fully

concealing gonocoxae and bases of telopodites. Gonopod coxites medium-sized, subcylindrical, fused

at base to a small membranous sternal remnant, poorly setose distodorsally, including a pair of very

closely placed, distalmost, particularly long setae. Cannulae slender, without peculiarities (Fig. 3A-B).

Telopodites in situ directed forward, held subparallel to each other, suberect, not crossing each other

mesally. Prefemoral (= densely setose) portion (pf) subcylindrical, very long and slender, untwisted,

almost not enlarged towards an oblique apical cingulum (c) delimiting it from femorite, with a long,

slightly curved, distodorsal process (pfp). Femorite clearly twisted, lamellar, about half as long as

pf, forming a large, rounded, apical lobe = solenophore (sph) to protect a smaller, parabasal lobe (lo)

carrying terminal part of seminal groove on its distal rib. Seminal groove first running fully on mesal

side of pf, then moving laterad above c onto distal rib of lo to end at tip of lo (Fig. 3C-E).

Remark

This new genus and species extends the range of Campodesmidae to Cameroon in the east, the family

also being recorded in that country for the first time.

Genus Campodesmus Cook, 1896

Campodesmus alobatus sp. nov.

urn:lsid:zoobank.org:act:5483AFA3-1222-4B5F-B7C7-B85EEE2FFB58

Figs 4-6

Diagnosis

Differs from other members of the genus in the complete lack of a parabasal process/lobe on the gonopod

telopodite which is not directed mesally, but held subparallel to the main body axis, coupled with much

more numerous, nearly undifferentiated and low tubercles on metaterga and the presence of deeply

incised and clearly separated antero- and caudolateral lobulations on the postcollum paranota.

Etymology

The name of this species emphasizes the absence of a dorsal/lateral parabasal process/lobe from the

gonopod telopodite.

VANDENSPIEGEL D. et al.. Two new Campodesmidae

Material examined

Holotype

IVORY COAST: 3, Appouesso, N 06°35’, W 003°28’, Foret Classee Bossematie, station 5A, pitfall

trapping, 30 Nov. 1994, leg. R. Jocque & N. Seabe, MRAC 17.264.

Paratypes

IVORY COAST: 1 3 , same locality, station 1A, 31 Oct. 1994, MRAC 17.272; 1 3 , same locality, station

5B, 30 Oct. 1994, MRAC 17.276; 1 3 (SEM) and 1 3 , same locality, 31 Oct. 1994, MRAC 17.278; 1 3 ,

same locality, station 5D, 31 Oct. 1994, ZMUM p2665. All leg. R. Jocque & N. Seabe.

Description

Length of holotype c. 17 mm, width on midbody pro- and metazonae 1.7 and 3.6 mm, respectively.

Paratypes (males) also c. 17 mm long, width on midbody pro- and metazonae 1.6-1.7 and 3.0 mm,

respectively. Coloration from light grey-brown to brown (Fig. 4); clypeolabral region, antennae,

prozonae, venter, legs and tip of epiproct pallid to light yellow; stricture as brown as metazonae.

Fig. 4. Habitus of Campodesmus alobatus sp. nov., 3 , holotype, lateral view. Photo by J. Brecko. Scale

bar =1.0 mm

European Journal of Taxonomy 139: 1-11 (2015)

Fig. 5. SEM micrographs of Campodesmus alobatus sp. nov., paratype, MRAC 17.278. A, D, G.

Anterior part of body, lateral, dorsal and ventral views, respectively. B, E, H. Midbody segments, lateral,

dorsal and ventral views, respectively. C, F, I. Posterior part of body, lateral, dorsal and ventral views,

respectively. J. Cross-section of a midbody segment, caudal view. K. Midbody paratergum, lateral view.

L. Fine structure of pro- and metazonae, as well as of stricture between them, subdorsal view. M.

Limbus and adjacent structures. Scale bars: A-J = 0.5 mm; K-L = 0.1 mm; M = 0.01 mm.

VANDENSPIEGEL D. et al.. Two new Campodesmidae

Head. Densely setose, clypeolabral region smooth, remaining parts densely microdenticulate-

microgranulate, interantennal isthmus c. 1.5 times as broad as diameter of antennal socket. Antennae

short and strongly clavate, C-shaped, antennomere 5 largest and longest, but only a little larger than 6 th ,

both with a dorso-apical group of tiny bacilliform sensilla; apical segment with usual four sensory cones

(Figs 4, 5G).

Body. With 20 segments (male). In width, head « collum = segment 2 < 3 < 4 < 5-17; thereafter

body rapidly tapering towards telson. Collum transversely ellipsoid, subflabellate, covering most of head

from above, dorsal surface rather regularly and densely tuberculate, with about 4-5 transverse rows of

round, setigerous, undifferentiated tubercles lying between a moderately crenulate/lobulate and regularly

rounded anterior margin and a clearly ribbed and almost straight caudal edge; small, but discernible

caudal paranota with a rather sharp lateral tooth (Figs 4, 5A, D, G). Prozonae smooth and poorly shining,

stricture finely microgranulate; metazonae dull, densely tuberculate and microgranulate over entire

surface; cerotegument well-developed, sometimes in places clothed with a crust of soil; dorsal surface

usually with two regular, mixostictic (= not regularly longitudinal) transverse rows of normally similar

round tubercles, much like on collum, lying between a fore row of denser and smaller knobs/tubercles

behind stricture and a densely ribbed caudal edge represented by a flatter row of oblong tubercules (Fig.

5A-F). Paranota 2 not enlarged compared to following ones (Figs 4, 5A). Postcollum paranota set rather

low (at about half of midbody height, thus dorsum being very strongly convex), strongly declivous,

continuing the outline of dorsum, but their tips not reaching level of venter (Fig. 5A-C, J); lateral margin

of paranota usually 5-lobulate, laterals continuing into even more deeply lobulate caudo- and anterolaterals

both at caudal and fore margin, respectively; caudo- and anterolateral lobulations continuing both caudal

and fore rows of tubercles at base of paranota, respectively (Figs 4, 5A-I). Caudal corner of paranota

increasingly strongly inclined caudad behind rear tergal margin only in segments 17-19 (Fig. 5C, F, I).

Tergal setae very short and thin (Fig. 5A-F). Ozopores inconspicuous, opening flush on tergal surface at

about midlength of paranota 5 and 7 off lateral margin. Pleurosternal carinae absent (Fig. 5G). Fimbus

microcrenulate, with microserrate tips of crenulations, these latter clearly separated from one another

(Fig. 5M). Epiproct short, small, subtruncate, slightly flattened dorsoventrally, tuberculate dorsally, with a

pair of rather large subapical papillae (Figs 4, 5C, F, I). Hypoproct nearly semi-circular with 1 + 1 caudal

setae very distinctly separated and borne on minute knobs (Fig. 51).

Fig. 6. Campodesmus alobatus sp. nov., paratype, $, MRAC 17.278. A-B. Feft gonopod, ventrolateral

and submesal views. C. Right gonopod, mesal view. Scale bars: A-B = 0.1 mm; C = 0.2 mm. Abbreviations

are explained in the text.

European Journal of Taxonomy 139: 1-11 (2015)

Sterna. Broad, almost twice as broad as coxa length, with a very deep axial impression (Fig. 5G-I).

Gonapophysis on male coxae 2 a small tubercle. Legs short, even in male about 0.9-1.0 times as long

as midbody height, very slender; in length, femur > tarsus > prefemur > postfemur = tibia = coxa; claw

very small, very slightly curved; ventral surface of tarsi densely setose, but forming no brushes.

Gonopod. Aperture transversely ovoid, large, its lateral and posterior edges slightly elevated, fully

concealing gonocoxae and bases of telopodites. Gonopod coxites medium-sized, subcylindrical, fused

at base to a small membranous sternal remnant, each with only one long distodorsal seta (Fig. 6A, B).

Cannulae long and slender, without peculiarities. Telopodites rather regularly curved caudad, in situ

held subparallel to each other, subcircular, long, slender and quite simple (Fig. 6). Prefemoral (= densely

setose) portion (pf) subcylindrical, rather long, totally devoid of outgrowths. Femorite (fe) with a small

parabasal denticle (d) on ventral face and a series of inconspicuous folds (f) on opposite side. Seminal

groove (sg) running entirely on mesal side, terminating at about halfway along fe, at base of a small

denticle (o), with neither a hairpad nor an accessory seminal chamber. Tip of fe slightly broadened and

bifid (Fig. 6C).

Remarks

The species described above is the 8 th known species of Campodesmus which, despite the peculiarities

referred to in the diagnosis, remains a rather typical congener. This is primarily evidenced by the

gonopod conformation which shows long and subcircular telopodites. Such non-gonopod characters

as low paranota, a reduced pore formula, short antennae and legs etc. also attest to the unequivocal

assignment of this new species to Campodesmus (cf. Hoffman 1951; Schiotz 1966; Demange 1971).

The clearly abbreviated seminal groove in C. alobatus sp. nov., which only extends to about the

midlength of an untwisted gonopod femorite, invites comparisons with other congeners. Regrettably,

no such information is available in the literature. Schiotz (1966) attempted to group most of the species

of Campodesmus , but more data are needed, including the course of the seminal groove, to reach more

secure conclusions.

Acknowledgements

The last two authors are greatly obliged to the Musee Royal de l’Afrique Centrale, Tervuren, Belgium

for the invitation to join this project. Special thanks go to Jonathan Brecko (MRAC) for taking the

colour pictures and to Christophe Allard for technical assistance.

References

Demange J.M. 1971. Myriapodes Diplopodes. Le massif des Monts Loma (Sierra Leone). Fascicule I.

Memoires de l Tnstitut Fondamental dAfrique Noire 86: 191-240.

Golovatch S.I. 2003. A review of the volvatory Polydesmida, with special reference to the patterns of

volvation (Diplopoda). African Invertebrates 44 (1): 39-60.

Hoffman R.L. 1951. The diplopod family Campodesmidae (Polydesmida). Journal of the Washington

Academy of Sciences 41 (6): 209-212.

Hoffman R.L. 1980. Classification of the Diplopoda. Museum d’histoire naturelle, Geneve.

Hoffman R.L. 1982. Diplopoda. In: Parker S.P (ed.) Synopsis and Classification of Living Organisms.

Vol. 2: 689-724. McGraw-Hill, New York.

Schiotz V. 1966. Notes on West African diplopods I. The family Campodesmidae (Polydesmida).

Nidensksbelige Meddelelser fra Dansk naturhistorisk Forening 129: 81-104.