Zoosphaerium coquerelianum (de Saussure & Zehntner, 1897 )

Wesener, Thomas & Wägele, Johann-Wolfgang, 2008, The giant pill-millipedes of Madagascar: revision of the genus Zoosphaerium (Myriapoda, Diplopoda, Sphaerotheriida), Zoosystema 30 (1), pp. 5-85 : 38-42

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https://doi.org/ 10.5281/zenodo.5391729

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Zoosphaerium coquerelianum (de Saussure & Zehntner, 1897 )


Zoosphaerium coquerelianum (de Saussure & Zehntner, 1897) View in CoL

( Figs 20-22 View FIG View FIG View FIG )

Sphaerotherium coquerelianum de Saussure & Zehntner, 1897 : pl. 1, figs 1c, 2a-c, pl., fig. 11; 1902: 42, pl. 1, fig. 1.

Heligmasoma errans Chamberlin, 1921: 8 View in CoL , n. syn.

Zoosphaerium amittum Chamberlin, 1921: 9 View in CoL , n. syn.

Zoosphaerium coquerelianum View in CoL – Jeekel 1999: 11 (lists species name). — Enghoff 2003: 20 (lists species name).

TYPE MATERIAL. — Zoosphaerium coquerelianum: VI. 18 2, leg. Coquerel, ♂ lectotype (broken, here designated), ( MNHN CB01 View Materials ) ; 2 ♀♀ paralectotypes ( MNHN CB01 View Materials ) ; Madagascar, 18 2, leg. Coquerel, no., 1 ♀ paralectotype ( MNHN CB018 View Materials ) .

Heligmasoma errans : ♂ holotype, no collection data ( MCZ 4 View Materials 92) ; 1 ♂, 2 ♀♀ paratypes ( MCZ 4 View Materials 93) .

Zoosphaerium amittum : ♀ holotype (MCZ 4709).

TYPE LOCALITY. — Zoosphaerium coquerelianum : Madagascar.

Heligmasoma errans : unknown.

Zoosphaerium amittum : uncertain, but probably Madagascar (“Whyman coll.”).

OTHER MATERIAL EXAMINED. — Madagascar. Leg. Grandidier, 2 ♂♂, 1 ♀ (MNHN CB017).

DIAGNOSIS. — Up to 3 mm long. Colour brown, posterior margin of tergites dark brownish, some smaller specimens with olive-greenish spots. Surface of tergites structured similar to an orange surface, glabrous. Anal shield well-rounded, glabrous. Apical antennomere with 18-29 sensory cones. Third joint of posterior telopods broad, with well-rounded tip. Posterior side with up to 7 sclerotized black teeth, 1 small non-sclerotized lobe and 3 sclerotized spines ( Fig. 20J, K View FIG ). Second joint with long and slim, curved process, basally with 1 large spine. One stridulation rib on male harp ( Fig. 20F View FIG ) and 2 or 3 on each side of female washboard. Anal shield with 2 locking carinae, first small, second more than 2. times longer than first ( Fig. 20B View FIG ). Operculum of vulva small, not protruding above coxa, apically with 2 well-rounded lobes protruding to same height. Interior plate of vulva protruding as high as operculum ( Fig. 21A, B View FIG ).


All species of the coquerelianum -group show similarly shaped telopods and opercula. Zoosphaerium coquerelianum is the only described species in this group that possess a single stridulation rib on the male harp. This feature identifies the large species unambiguously.


Body measurements: males (3 specimens): length up to 22.00 (largest broken), width of thoracic shield up to 18., height of thoracic shield up to 10.. Females (specimens): length up to 2.4, width up to 32.4, height up to 1..

Habitus: body massive and high ( Fig. 20A View FIG ). Tergites and anal shield without hair, but surface completely covered with irregular small pits, comparable to an orange surface.

Coloration: faded in alcohol. Tergites brown, posterior margin with thin dark brown line. Some specimens with olive green spots, resembling possibly the remains of original coloration. Head, antennae and tips of legs dark green.

Head: posterior margin of head towards collum without small hair.

Antennae: long, sixth joint of cylindrical shape with 18-29 sensory cones. First joint remarkably



broader than others. At least first joint with sclerotized teeth. Mouthparts not dissected.

Collum: anterior and posterior margin with some long hairs arranged in one row. Remaining part of collum with some long, isolated hairs.

Thoracic shield: thoracic shield surface structured like those of tergites ( Fig. 20A View FIG ).

Tergites: anterior paratergite depressions of tergites 3-12 with isolated hairs.Tips of paratergites margins projecting posteriorly.

Endotergum: internal section with short spines and isolated bristles. One row of large circular cuticular patterns located between marginal ridge and internal area. Marginal brim strong and straight ( Fig. 22A, B View FIG ). Externally 2 rows of marginal bristles, standing closely to one another. Bristles scaly ( Fig. 22C, D View FIG ), protruding up to 1/2 towards area between basal point of bristles and tergite margin ( Fig. 22E View FIG ).

Anal shield: well-rounded, neither bell-shaped nor tapered ( Fig. 20A View FIG ). Like tergites completely covered with numerous small irregular pits, surface orangelike. Underside carrying two well-developed black locking carinae on both sides, anterior one small, but well-developed, projecting posteriorly strongly towards anal shield margin. Posterior carina more than 2. times longer than first, straight ( Fig. 20B View FIG ). Locking carinae separated by a distance equal to half of shorter carina length ( Fig. 20B View FIG ).

Legs: first tarsi with or, second with -8 long ventral spines. Third tarsi with 7-9 long ventral spines. First two tarsi with only weakly curved claws and without apical spine. Tarsi of legs 3-21 with curved claws, 8 or 9 ventral spines and an apical spine. Ventral spines of legs dispersed through whole inner side of tarsus. In 9th leg femur 1.9 times longer than wide, tarsi 4.9 times longer than wide. Femora with crenulated ridge of medium length ( Fig. 20D View FIG ).

Stigma-carrying plate: first stigma-carrying plate lobe short and well-rounded, protruding up to apical edge of coxa 1. Lobe curved towards coxa, its broad, well-rounded tip thinner than basal part ( Fig. 20C View FIG ).

Female sexual characters: second leg pair on lateral side with some small, black triangular spines ( Fig. 21A, B View FIG ). Subanal plate with a washboard, consisting of well-developed stridulation ribs, 2 up to 3 ribs on each side. Stridulation ribs symmetrical, sometimes non-continuous, especially the inner rib. Ribs long, ending just before posterior margin. Vulva large, covering slightly more than 1/2 of coxa. Operculum low, not surmounting coxa. Apical margin of operculum medially notched, with two rounded lateral tips, both of same height ( Fig. 21A, B View FIG ). Inner plate long and broad, protruding as high as operculum ( Fig. 21A, B View FIG ).

Male sexual characters: male gonopore covered with single, large, sclerotized, undivided and rounded plate. Apical part of plate membranous. Gonopore covering 1/3 of height and a little less than 1/2 of coxal width ( Fig. 20D View FIG ). Anal shield similar to those of females.

Anterior telopods: first joint with a stridulation harp and one stridulation rib. Rib straight, broad and well-developed ( Fig. 20F View FIG ). Second joint on posterior side with lobe-like, weakly curved process with rounded edges ( Fig. 20 View FIG G-I). Point of process protruding almost as high as third joint, towards third joint apically with sclerotized spots and basally with single spine ( Fig. 20H View FIG ). Third joint with strong tip bending inwards, protruding twice as high as second joint process. Joint with a cavity juxtaposed to second joint process. Apical margin of third joint well-rounded, at internal margin towards cavity with numerous sclerotized spots and two small spines, a third spine located closely towards tip ( Fig. 20 View FIG G-I).

Posterior telopods: third joint weakly curved, becoming only slightly slimmer towards tip. Hollowed-out margin towards second joint with 1 small non-sclerotized lobe and up to 3 smaller, sclerotized spines, one of them located apically, the two other ones at half of its length ( Fig. 20J View FIG ). Third joint on posterior aspect with 1 row of c. 7 sclerotized black teeth ( Fig. 20K View FIG ). Process of second joint slim, almost as long as third joint, with a weakly curved tip. Anterior side basally with 1 sclerotized spine. Third joint on posterior side with few isolated hairs, on rest of surface hairs absent. First and second joints on both sides, particularly on posterior side, with numerous hairs ( Fig. 20J, K View FIG ). Inner horns of syncoxite apically curved posteriorly.Tips sharp, whole horns covered with numerous small hairs.


The material contains too few specimens to evaluate the intraspecific variation. The vulvae of small and gigantic specimens feature the same pattern ( Fig. 21A, B View FIG ), different to the drawing given in the first description (de Saussure & Zehntner 1897).


No precise locality data exist for this species. No material was found in more recently collected material. This species is only known from the type series.


It is unusual that no material of such a large species as Z. coquerelianum was found in the Museum collections. It could be suggested that this species is extinct.


All type specimens of Heligmasoma errans are similar to the types of Z. coquerelianum . Chamberlin (1921) obviously did not compare H. errans with any described Malagasy Sphaerotheriida species when he erected the new genus Heligmasoma . The same fits to Zoosphaerium amittum which was described by Chamberlin (1921) in the same paper. Chamberlin mentioned in the first description of both species only general characters, which seem to fit on almost all known Sphaerotheriida species. Hoffman (1980: 2) already noted to the genera described by Chamberlin that “although it seems probable that both are junior synonyms of some common and well-known genus”. However, if the coquerelianum species-group will later be elevated to genus rank, the genus name Heligmasoma would be available for this group.


Museum National d'Histoire Naturelle














Zoosphaerium coquerelianum (de Saussure & Zehntner, 1897 )

Wesener, Thomas & Wägele, Johann-Wolfgang 2008

Zoosphaerium coquerelianum

ENGHOFF H. 2003: 20
JEEKEL C. A. W. 1999: 11

Heligmasoma errans

CHAMBERLIN R. V. 1921: 8

Zoosphaerium amittum

CHAMBERLIN R. V. 1921: 9
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