Zoosphaerium coquerelianum (de Saussure & Zehntner, 1897)

Wesener, Thomas & Sagorny, Christina, 2021, Seven new giant pill-millipede species and numerous new records of the genus Zoosphaerium from Madagascar (Diplopoda, Sphaerotheriida, Arthrosphaeridae), European Journal of Taxonomy 758 (1), pp. 1-48 : 18-20

Zoosphaerium coquerelianum species-group

Remarks

Species can be identified as members of the Zoosphaerium coquerelianum species-group based on morphological characters of the posterior telopods ( Wesener & Wägele 2008; Wesener 2009). Telopoditomere 3 only carries small sclerotized teeth and is always slightly longer and>1.5 times as wide as the immovable finger. Similar telopod characters occur in some members of the Z. platylabum species-group, but the locking carinae in this group are located close to the last pleurites, while they are positioned close to the anal shield margin in species of the Z. coquerelianum group. The speciesgroup now contains 23 described species distributed in humid to subhumid forests all over Madagascar. Species of this group are conspicuously absent from the spiny forest ecosystem and the western dry forest (Wesener 2009; Wesener et al. 2010b; Sagorny & Wesener 2017; Wesener & Anilkumar 2020).

Key to the species of the Zoosphaerium coquerelianum group

1. Male harp with two to three stridulation ribs..................................................................................2

– Male harp with a single stridulation rib........................................................................................11

2. Tergites glossy, anteriorly shiny light green, posteriorly dark green. Specimen more than 50 mm long. Anal shield not well-rounded, with steep slope. Movable finger of posterior telopods 1.6 times as long as wide, apically with groove ...................................................... Z. amabile Wesener, 2009

– Coloration pattern different, or if similar then specimens less than 35 mm long. Anal shield wellrounded or bell-shaped. Movable finger of posterior telopods 1.9–2.5 times as long as wide, without apical groove.........................................................................................................................3

3. Antenna with four apical cones.......................................................................................................4

– Antenna with>10 apical cones......................................................................................................5

4. Anal shield surface covered with short hairs. Posterior locking carina of anal shield 2.5–3 times as long as anterior carina ............................................. Z. alluaudi (de Saussure & Zehntner, 1902)

– Anal shield surface glabrous. Posterior locking carina of anal shield more than 3.5 times as long as anterior carina ............................................. Z. voeltzkowianum (de Saussure & Zehntner, 1897)

5. Male harp with three stridulation ribs. Endotergum with three rows of marginal bristles, bristles slightly extending beyond tergite. Sclerotized teeth present on antennomeres 1–5. Movable finger of posterior telopods 2 times as long as wide. Posterior locking carina of anal shield four times as long as anterior carina. Male anal shield bell-shaped ....... Z. trichordum Wesener, 2009

– Male harp with two stridulation ribs. Endotergum with one up to two rows of marginal bristles, bristles not reaching tergite margin. Sclerotized teeth only present up to antennomere 3 or 4......6

6. Only up to eight ventral spines on walking legs.............................................................................7

– At least nine ventral spines on walking legs.................................................................................8

7. Movable finger of posterior telopod thick, 2.5 times as long as wide. Leg 3 with an apical spine. On anal shield, second locking carina 2–3 times as long as first ............ Z. ambatovaky sp. nov.

– Movable finger of posterior telopod very thick, 2 times as long as wide. Leg 3 without an apical spine. On anal shield, second locking carina 4–5 times as long as first .............................................. ......................................................................................... Z. muscorum Wesener & Bespalova, 2010

8. Male anal shield well-rounded, with slight bell shape. Posterior locking carina of anal shield six times as long as anterior carina. Up to 10 ventral spines on walking legs .............................. .................................................................................... Z. bambusoides Wesener & Bespalova, 2010

– Male anal shield well-rounded. Posterior locking carina of anal shield three to four times as long as anterior carina. Usually more than 10 ventral spines on walking legs.............................9

9. Leg 3 without an apical spine. Specimens only slightly more than 30 mm long. Tergites jadelike green with thin brown posterior margin, texture glossy. Operculum of vulva deeply notched, strongly extending beyond coxa. Apical disc with ca 14 apical cones ............................................... .......................................................................................................... Z. smaragdinum Wesener, 2009

– Leg 3 with an apical spine. Specimens more than 50 mm long. Tergites never jade-like, not glossy. Operculum of vulva not protruding above coxal margin. Apical disc with more than 20 apical cones................................................................................................................................................10

10. Specimens more than 60 mm long. Tergites dark-green, covered with numerous large pits and short setae. Sclerotized teeth present on antennomeres 1–3, disc with ca 45 apical cones. Endotergum with two rows of marginal bristles ......... Z. villosum Wesener & Sierwald, 2005

– Specimens less than 60 mm long. Tergites light-brown, glabrous. Sclerotized teeth present on antennomeres 1–4, disc with 20–30 apical cones. Endotergum with a single row of marginal bristles .............................................................................. Z. bemanevika Sagorny & Wesener, 2017

11. Movable finger 3 times as long as wide ................................................ Z. ambrense Wesener, 2009

– Movable finger <2.5 times as long as wide....................................................................................12

12. Antenna with four apical cones. Movable finger of posterior telopods at inner margin with two membranous lobes...........................................................................................................................13

– Antenna with>10 apical cones. Movable finger of posterior telopods at inner margin with a single membranous lobe............................................................................................................................17

13. Process of second podomere of anterior telopod not visible in anterior view. Fifth antennomere with a field of sensilla basiconica. Collum with isolated, long setae. Endotergum with one row of large cuticular impressions and a second row of much smaller impressions; bristles long, strongly protruding above tergite ........................................................... Z. bilobum Wesener, 2009

– Process of second podomere of anterior telopod visible in anterior view.................................14

14. Podomere 3 of anterior telopod with crenulated teeth...................................................................15

– Podomere 3 of anterior telopod without crenulated teeth.............................................................16

15. Collum glabrous, with a few setae at corners on either side of head. Endotergum with two rows of regularly distributed circular impressions, marginal bristles strongly protruding above tergite. Sensilla basiconica present on antennomeres 1 and 2. Anal shield weakly bellshaped ................................................................ Z. trigrioculatum Wesener & Bespalova, 2010

– Collum glabrous, anterior margin with two rows of setae. Endotergum with a single row of elliptical cuticular impression, marginal bristles protruding to margin. Sensilla basiconica absent. Anal shield well rounded .................................. Z. bartolozzi Anilkumar & Wesener, 2020

16. Collum glabrous. Endotergum with a single row of large cuticular impressions, marginal bristles slightly protruding above tergite margin. Second leg with 4 or 5 ventral spines. Anal shield tapering ........................................................................................................................ Z. isalo Wesener, 2009

– Collum glabrous. Endotergum with a single row of slightly rounded elliptical cuticular impressions, marginal bristles protruding to margin. Second leg with 6 or 7 ventral spines. Anal shield well rounded .................................................................. Z. taiti Anilkumar & Wesener, 2020

17. First pair of stigmatic plates greatly enlarged, with a well-rounded tip......................................18

– First pair of stigmatic plates triangular, tapering towards tip.......................................................19

18. Operculum of vulva ending clearly before coxal margin, deeply notched, divided into two large, well-rounded, similar-sized lobes. Third podomere of anterior telopod laterally with ca five large sclerotized teeth. Sensilla basiconica on first antennomere absent. Posterior locking carina of anal shield two times as long as anterior carina.Endotergal marginal bristles protruding up to tergite margin. Movable finger of posterior telopods thick, 1.9 times as long as wide ...... Z. tampolo Wesener, 2009

– Operculum of vulva ending before coxal margin, elongated into a single median lobe, lobe only shallowly notched at tip. Third podomere of anterior telopod without any large sclerotized teeth. Sensilla basiconica on first antennomere present. Locking carinae of anal shield of equal length. Endotergal marginal bristles very short, ending far before tergite margin. Movable finger of posterior telopods thick, 2.1 times as long as wide ............... Z. ignotum Wesener, 2009

19. Tergite texture orange-peel like. Collum covered with isolated, long hairs. Body size of adult female> 50 mm. Endotergum with two rows of marginal bristles and a single row of large circular cuticular impressions........................................................................................................20

– Tergite texture glossy or smooth. Collum glabrous. Body size of adults <35 mm. Endotergum with a single row of marginal bristles and either without or with two rows of cuticular impressions.........................................................................................................................................22

20. Posterior locking carina on anal shield 2.5 times as long as anterior carina. Third podomere of anterior telopods without large sclerotized teeth. Anal shield glabrous ............................................ ........................................................................... Z. coquerelianum (de Saussure & Zehntner, 1897)

– Posterior locking carina on anal shield>4 times as long as anterior carina. Third podomere of anterior telopods with at least three large sclerotized teeth. Anal shield covered with short hair.............................................................................................................................................21

21. Third podomere on anterior telopods with 3 or 4 crenulated teeth. Endotergal marginal bristles very short, protruding up to ⅓ of distance to tergite margin. Tips of vulvar operculum widely separated ................................................................................................. Z. mangabe Wesener, 2020

– Third podomere on anterior telopods with ca seven large sclerotized teeth. Endotergal marginal bristles protruding up to tergite margin. Tips of vulvar operculum fused ...................................... ............................................................................................................. Z. tainkintana Wesener, 2009

22. Tergites glossy. Female anal shield bell-shaped, in male with steep slope. Fixed finger of posterior telopods wide, only slightly curved towards movable finger. Movable finger 1.8 times as long as wide, on posterior side with ten small sclerotized teeth. Sclerotized teeth on antenna only present on first antennomere. Endotergum without cuticular impressions ......... Z. viridissimum Wesener, 2009

– Tergites matte. Anal shield well-rounded. Fixed finger of posterior telopods slender, strongly curved towards movable finger. Movable finger 2.4 times as long as wide, on posterior side with ca 25 small sclerotized teeth. Sclerotized teeth present on antennomeres 1–3. Endotergum with a single row of large and a second row of smaller circular cuticular impressions ............... ......................................................................................................... Z. denticulatum Wesener, 2009