Zoosphaerium arborealis, Wesener & Sierwald, 2005
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Zoosphaerium arborealis sp. nov.
Paratypes: 2 m., 8 f. (mature) FMNH ; 1 m., 2 f. (mature) CAS ; 1 m., 2 f. (mature) ZSMC ; 3 f. SMNG VNR 13554 View Materials ; Madagascar; Province Toliara; Mandena ; littoral forest; in leaf litter and on trees; 24°56.51’S, 046°59.70’E; IV.2003; leg. T. Wesener. GoogleMaps 2 m., 2 f. FMNH . Madagascar; Province Toliara; Sainte Luce ; littoral forest; in leaf litter; 24°46.77’S, 047°10.28’E; 07.IV.2003; leg. T. Wesener. GoogleMaps
Other Material: 29 f. (mature) Université de Antananarivo. Madagascar; Province Toliara; Mandena ; littoral forest; in leaf litter; on trees; many inside of dead stems of Pandanus sp. ; 24°56.51’S, 046°59.70’E; IV.2003; leg. T. Wesener. GoogleMaps 7 m., 11 w. (immature), FMNH. Madagascar; Province Toliara; Mandena ; littoral forest; in leaf litter; on trees; many inside of dead stems of Pandanus sp. ; 24°56.51’S, 046°59.70’E; IV.2003; leg. T. Wesener. GoogleMaps 1 m., 2 w. (immature), FMNH. Madagascar; Province Toliara; Sainte Luce ; littoral forest; in leaf litter;; 24°46.77’S, 047°10.28’E; 07.IV.2003; leg. T. Wesener. GoogleMaps 3 m., 13 w. (mature) FMNH. Locality: Province Fianarantsoa; Parc National de Andringitra , Forêt d'Ravaro , 12.5 km SW Antanifotsy, 22°12.7’S, 46°50.7’E, 1500 m, 9–16.i.2000, undisturbed midelevation forest /w Acanthaceae , leg. S. Goodman. GoogleMaps
Diagnosis: Mediumsized pillmillipede up to 40 mm in body length. Males distinctly smaller than females. Tergites brown, legs and antennae red ( Fig. 29a View FIGURES 29 ). Body smooth or with a few hairs, anal shield slightly tapered ( Fig. 19a View FIGURES19 ). Antennae with 7–24 sensory cones ( Fig. 22b View FIGURES 22 ), first and second joint with invagination, all joints at least basal with sclerotized teeth ( Fig. 22a View FIGURES 22 ). First sternite at inner margin straight ( Fig.19d View FIGURES19 ). Anal shield ventrally on each side with two welldeveloped black locking carinae. Anterior locking carinae short, posterior one 3–4 times longer than anterior carina ( Fig. 19c View FIGURES19 ). Anterior telopods with two stridulation ribs, number of ribs on female washboard variable ( Figs. 20b View FIGURES20 , 21a View FIGURES 21 ).
Similar species: Z. arborealis is closely related with Z. lambertoni ( Brolemann, 1922) . Nevertheless, Z. arborealis the inner horns and inner lobes of the posterior telopods are unique in shape ( Figs 21e, f View FIGURES 21 ). Additionally, the locking carinae of the anal shield are strongly developed in Z. arborealis , but weakly developed in Z. lambertoni ( Brolemann, 1922: 235–236, " font défaut en tant quíarêtes pigmentées [...]"). The type specimen of Z. lambertoni ( Brolemann, 1922) , deposited at the L`Académie Malgache in Antananarivo / Madagascar, cannot be located. Thus, the species cannot be compared to Z. arborealis . All other species of Zoosphaerium with similar posterior telopods like Z. anomalum ( DeSaussure & Zehntner, 1897) have a differently formed anal shield. The lower margin at the posterior end of the anal shield is never tapered in Z. arborealis .
Description: Males (14 specimens): length: 19.0–26.45, width of thoracic shield: 9.65–13.15, height of thoracic shield 5.5–7.2. Females (45 specimens): length: 21.0–40.0, width: 10.75–20.0, height: 6.75–11.35.
Habitus: tergites more posterior higher than anterior ( Fig. 19a View FIGURES19 ). Tergites very smooth, sometimes with a broad patch of hairs at the anterior and posterior margin.
Coloration: tergites brown, posterior margin darker ( Figs. 29a–b View FIGURES 29 ), smaller specimen olivebrown. Antennae and legs darkorange to red. Red color fades in alcohol.
Head: with numerous hairs and hairpits especially around the labrum and lateral of eyes. A few single, long hairs around the eyes and distributed on the entire head. Posterior margin of head towards the collum with a patch of short hairs ( Fig. 23d View FIGURES 23 ). Edges of antennal groove with 5–6 small spines and sclerotized teeth ( Fig. 23b View FIGURES 23 ). Head with up to 90 ocelli on each side ( Fig. 23c View FIGURES 23 ). Tömösváry organ inside with a corallike structure ( Fig. 23e View FIGURES 23 ).
Antennae: shaped as described for the genus. Length of antennomeres: 1>2>3=4=5<6; sixth antennomere longest, of cylindrical shape ( Figs. 22a View FIGURES 22 ) with 7–24 sensory cones ( Fig. 22b View FIGURES 22 ). First antennomere much broader than the others ( Fig. 22c View FIGURES 22 ). Sclerotized teeth at the base of antennomeres one through six, reaching the apical border at first antennomere. First and second joint with invaginations ( Fig. 22c View FIGURES 22 ).
Mandible: with seven rows of pectinate lamellae; number of teeth declining proximally ( Fig. 27b View FIGURES 27 ). Condylus with two strongly developed steps near the apical margin ( Fig. 27c View FIGURES 27 ). Gnathochilarium: ventral side with bristles, only few on the lingual lamella ( Fig. 24a View FIGURES 24 ). With a field with three sensory cones lateral to the palpi ( Fig. 24b View FIGURES 24 ). Two different types of sensory cells on the central pads: long ones without a pit and smaller, cylindershaped ones with a central pit ( Fig. 25a View FIGURES 25 ). Sensory cones on palpi distributed regularly over the tip ( Fig. 25b View FIGURES 25 ). Epipharynx: shaped as in other members of this genus ( Figs. 26b View FIGURES 26 , 27a View FIGURES 27 ).
Collum: Anterior margin bald, in the middle with a row of very small, darkbrown sclerotized teeth. Posterior margin on each half with 4–6 hairs, inserting lateral and at the edges. Rest of collum hairless. Thoracic shield: A few hairs in the concave lateral extension of the thoracic shield, especially near the margins. Anterior marginal brim broader than the rest of the brim ( Fig. 19a View FIGURES19 ).
Anterior paratergite depression of tergites 3–12 with many hairs, on the anterior margin with a few ribs ( Fig. 19a View FIGURES19 ), otherwise tergites are almost hairless and smooth. Tips of the posterior margins of paratergites 8–12 projecting posteriorly.
Endotergum with two or more rows of marginal bristles. Internal part with short spines and few, isolated bristles. Bristles scaly ( Fig. 25c View FIGURES 25 ). Marginal ridge straight. Interior of marginal ridge one row of round flat nodules ( Fig. 26a View FIGURES 26 ).
Anal shield: lower margin at posterior end tapered ( Fig. 19a View FIGURES19 ). Only in some males near posterior margin with many short (sensory?) hairs. Ventral side carries two black locking carinae on both sides, anterior one similar to those of tergites, posterior one more than three and a half times longer than anterior one, at posterior end nearer to the margin than anteriorly. Locking carinae separated from each other by a distance equal to the double length of the shorter carina. Distinct suture present between both carinae. Where the suture reaches the margin of the anal shield, a small, distinct triangular invagination is visible ( Fig. 19c View FIGURES19 ).
Legs: tarsi of first pair of legs with 4–6, those of second with 6–9 ventral spines, weakly curved claws and without apical spine. Tarsi of legs 3–21 with curved claws, 10–14 ventral spines and one apical spine. 9th leg pair without coxal lobe but with a few small, black triangular spines ( Fig. 19b View FIGURES19 ). Coxae of all legs at the inside margin covered with many long hairs, the following leg joints with a few isolated, long hairs at the inner margin. Femur with a dark crenulated ridge ( Fig. 19b View FIGURES19 ).
Sternite: first sternite lobe broad and short, reaching the apical edge of the coxae of first leg pair, covered sparsely with hairs ( Fig. 19d View FIGURES19 ). Apical margin rounded regularly and covered with many long hairs. Basal margin with few hairs ( Fig. 19d View FIGURES19 ). Inner margin of sternite remarkably straight. Sternites three and beyond with a spine like process which reaches about the stigma opening of the anterior sternite.
Female sexual characters: second leg pair without coxal lobe, but on exterior margin with some short, triangular black spines ( Fig. 20a View FIGURES20 ). Vulvae welldeveloped, covering more than 2/3 of the coxa ( Fig. 20a View FIGURES20 ). Operculum small, ending before the apical edge of coxae. Apical margin of operculum constricted in the middle (subreniform) with two broadly rounded equally sized lateral tips. Both parts of operculum of same height. Basal margin almost straight. Exterior and inner plates (EP, IP) of vulva below the operculum surrounding the basal margin of the operculum. Inner plate very long and broad, reaching almost 2/3 of height of operculum. Exterior plate not as long as the inner one, reaching only the basal 1/2 part of the operculum. Cyphopod sclerites consisting of two triangular apical sclerites and a much larger third sclerite formed like a tuningfork; all visible as dark structures near the suture of the vulva between inner and exterior plate ( Fig. 20a View FIGURES20 ). Subanal plate with washboard, consisting of irregular, asymmetrical 1–4 stridulation ribs. Length of ribs very variable ( Fig. 20b View FIGURES20 ).
Male sexual characters: second leg pair without coxal lobe, but on exterior margin with some short, triangular black spines and covered with many hairs ( Fig. 21d View FIGURES 21 ). Male gonopore covered half with a sclerotized, undivided, rounded plate, which is slightly excavated at apical margin. Gonopore plate covered with hairs. Apical part of plate membranous ( Fig. 21d View FIGURES 21 ). Anal shield without any invagination or other sexual dimorphism, but in some cases with patch of (sensory?) hairs at posterior margin.
Anterior telopods: syncoxite of anterior telopods with few isolated hairs. First joint with a stridulation harp and two stridulation ribs. Inner ridge short, reaching only 1/2 of the length of the more laterally ridge. Lateral ridge long, beginning at the basal margin of the first joint and ending just at its apical margin ( Fig. 21a View FIGURES 21 ). Posterior side of the first joint inside with an impression. Margins of impression laterally with one row of hairs. First joint with exception of margins on both sides bold. Second joint on posterior side with a lobelike apically stout process ( Figs. 21b View FIGURES 21 ). Tip of process reaching only slightly above the third joint. Towards the third joint with sclerotized spots ( Fig. 21c View FIGURES 21 ). Third joint inner margin drawn out, at apical margin broad. Apical margin almost straight, only slightly excavated near the middle, tip of second joint process visible. Inner margin of third joint towards second joint process with some spines and sclerotized spots, juxtaposed to the spots on the second joint ( Fig. 21c View FIGURES 21 ). Third joint apically with some thick bristles ( Figs 21a–b View FIGURES 21 ).
Posterior telopods: movable finger (third joint) of chela long and slightly curved towards the immovable finger (process of second joint, Fig. 21e View FIGURES 21 ). Invagination towards the immovable process of the second joint with 4–5 small, nonsclerotized spines, after one third of length of third joint with one big spine, after half of length with one fingershaped process. Third joint on posterior side with 17 th sclerotized teeth, size of teeth declining basally ( Fig. 21f View FIGURES 21 ).
Immovable digit (process of second joint) almost straight with slightly curved tip. Anterior side basally with one spine, apically with numerous sclerotized spots juxtaposed the invagination of the third joint. Chelae only basally with hairs, on rest of surface hairs absent ( Figs. 21f View FIGURES 21 ). Inner horns (IH) of syncoxite with a pointed tip ( Figs 21e–f View FIGURES 21 ).
Distribution & Ecology: This species has an extraordinarily wide distribution. It is the only species described from littoral forests, which is not restricted to one forest site. Zoosphaerium arborealis was also discovered in another ecosystem, in the montane forest of PN Andringitra at over 1200 m elevation.
In the littoral forests this species was collected in the fragments of Mandena and Sainte Luce. In both forests Z. arborealis is associated with a species of the giant pillmillipede genus Sphaeromimus splendidus Wesener & Sierwald (in press) and it also occurs in Sainte Luce sympatrically with Zoosphaerium sp. Sainte Luce.
To date, all 33 species of Malagasy pill millipede genus Zoosphaerium were known from museum material only. Observation of living specimens are presented here for the first time. During an expedition in beginning of April 2003 to the Malagasy southern littoral forests, the first author observed the activity, diversity, behavior and habitats of Zoosphaerium arborealis in the littoral forest fragment near Mandena ( Fig. 30 View FIGURE30 ).
This species was common in the littoral forest of Mandena. On 18 collection days> 300 specimens were observed. Smaller individuals were often found in dead, rotten wood, especially in dead, excavated stems of Pandanus sp. ( Pandanaceae ), normally 5– 15 specimen of almost the same size in a single stem. These stems were hollow, filled with numerous fecal pellets, indicating that the species feeds on the soft and stringy contents of the stems. Numerous smaller specimens were discovered in piles of dead wood, which were common in this forest, especially near the tracks. In these piles of dead wood, the giant pillmillipedes occurred together with larvae of Scarabaeidae and Cestonidae ( Insecta : Coleoptera ) and spirostreptids ( Diplopoda ), the most common arthropods.
The leaf litter was thin in the littoral forest of Mandena, after 1–2 cm the soil was strongly riddled with fine roots of trees. Below 20 cm the soil is sandy. After a few dry days, the soil and root layer and leaf litter dry out. Apparently, the sandy ground cannot hold the water and the layer of leaf litter is too thin to prevent moisture loss of the layers below. Only a few animals could be detected in this dry leaf litter, in some places Blattodea ( Insecta ) were found.
Adult specimens of Z. arborealis were collected in such places or under wood during drier days. Numerous pellets show that the animals use these moist microhabitats as retreats and for feeding. In the moist patches some Isopoda and Amphipoda (both Crustacea) could be found as well.
Adults of Z. arborealis were collected at dawn after rainfall during the night and also after heavy rainfalls in the thin leaf litter. A day earlier, before the rain, no specimens were observed at these sites. Numerous specimens of all sizes could be observed after heavy rainfalls walking on the tracks. It must be noted that adult specimen were observed climbing tree trunks more than 150 cm high, feeding on the stem ( Fig. 29a View FIGURES 29 ). While climbing on trees, members of this species do not roll into a ball after mechanical disturbance, only the head is tucked under the thoracic shield. Such arboreal behavior was previously reported only from SouthAfrican giant pillmillipedes of the genus Sphaerotherium ( Haacker & Fuchs 1972) .
No eggs were found in dissected females. The smallest juvenile stadium had 19 leg pairs. No matings were observed.
Predation: Tergites of smaller specimen of Z. arborealis were found in a layer of arthropod remains around ant holes of a big, red ant species in Mandena. This layer contained mostly seeds of plants and body rings of "juliform"diplopods. Living rolledup specimens of Z. arborealis were placed near ants and ant holes, but the ants did not attack the millipedes. One big (– 35 mm body length) specimen of Z. arborealis was placed in a cage with a small mongoose (Viveriidae, Galidia elegans ) from the forest of Mandena. The rolledup pill millipede was detected immediately, broken with lateral teeth and eaten by the mongoose, except for the intestines and tergites. It is quite possible that Galidia elegans is a natural predator of pill millipedes. One female collected in the littoral forest of Sainte Luce had fungi on the antennae. Specimens of all three localities were strongly covered with mites (Arachnida: Acari), it is unknown if this mites are commensals or parasites.
Conservation: This species has a wide distribution. One of the sites where Z. arborealis was collected is already protected. Nevertheless, the distribution of this species is scattered due to the vast destruction of natural forests on Madagascar.
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